Sitticine jumping spiders: phylogeny, classification, and chromosomes (Araneae, Salticidae, Sitticini)

ZooKeys 925: 1-54 (2020) A peer-reviewed open-access journal
doi: 10.3897/zookeys.925.3969 | RESEARCH ARTICLE $Z00Ke y S

http:/ / ZOO keys -pen soft. net Launched to accelerate biodiversity research

Sitticine jumping spiders: phylogeny, classification, and
chromosomes (Araneae, Salticidae, Sitticini)

Wayne P. Maddison', David R. Maddison’,
Shahan Derkarabetian?*, Marshal Hedin?

| Departments of Zoology and Botany and Beaty Biodiversity Museum, University of British Columbia, 6270
University Boulevard, Vancouver, British Columbia, V6T 1Z4, Canada 2. Department of Integrative Biology,
Oregon State University, Corvallis, OR 97331, USA 3 Department of Biology, San Diego State University,
San Diego, CA 92182, USA 4 Department of Organismic and Evolutionary Biology, Harvard University,
Cambridge MA 02138, USA

Corresponding author: Wayne P. Maddison (wayne.maddison@ubc.ca)

Academic editor: J. Miller | Received 4 September 2019 | Accepted 5 February 2020 | Published 8 April 2020
http://zoobank. ore/BB966609-0878-49A 1-B13C-138C2495E6B7

Citation: Maddison WP, Maddison DR, Derkarabetian S$, Hedin M (2020) Sitticine jumping spiders: phylogeny,
classification, and chromosomes (Araneae, Salticidae, Sitticini). ZooKeys 925: 1-54. https://doi.org/10.3897/
zookeys.925.39691

Abstract

The systematics of sitticine jumping spiders is reviewed, with a focus on the Palearctic and Nearctic re-
gions, in order to revise their generic classification, clarify the species of one region (Canada), and study
their chromosomes. A genome-wide molecular phylogeny of 23 sitticine species, using more than 700
loci from the arachnid Ultra-Conserved Element (UCE) probeset, confirms the Neotropical origins of
sitticines, whose basal divergence separates the new subtribe Aillutticina (a group of five Neotropical
genera) from the subtribe Sitticina (five genera of Eurasia and the Americas). The phylogeny shows that
most Eurasian sitticines form a relatively recent and rapid radiation, which we unite into the genus At
tulus Simon, 1868, consisting of the subgenera Sitticus Simon, 1901 (seven described species), Attulus (41
described species), and Sittilong Prészynski, 2017 (one species). Five species of Aztulus occur natively in
North America, presumably through dispersals back from the Eurasian radiation, but an additional three
species were more recently introduced from Eurasia. Attus palustris Peckham & Peckham, 1883 is consid-
ered to be a full synonym of Euophrys floricola C. L. Koch, 1837 (not a distinct subspecies). Attus sylvestris
Emerton, 1891 is removed from synonymy and recognized as a senior synonym of Sitticus magnus Cham-
berlin & Ivie, 1944. Thus, the five native Attu/us in North America are Attulus floricola, A. sylvestris, A.
cutleri, A. striatus, and A. finschi. The other sitticines of Canada and the U.S.A. are placed in separate gen-
era, all of which arose from a Neotropical radiation including Jollas Simon, 1901 and Tomis FO.Pickard-
Cambridge, 1901: (1) Attinella Banks, 1905 (A. dorsata, A. concolor, A. juniperi), (2) Tomis (T; welchi), and

Copyright Wayne P. Maddison et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
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2 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

(3) Sittisax Proszytiski, 2017 (S. ranieri). All Neotropical and Caribbean “Sitticus” are transferred to either
Jollas (12 species total) or Tomis (14 species). Attinella (three species) and Tomis are both removed from
synonymy with Sitticus; the synonymy of Sitticus cabellensis Prészytski, 1971 with Pseudattulus kratoch-
vili Caporiacco, 1947 is restored; Pseudattulus Caporiacco, 1947 is synonymized with Tomis. Six generic
names are newly synonymized with Aztulus and one with Aftinella. Two Neotropical species are described
as new, Jollas cupreus sp. nov. and Tomis manabita sp. nov. Forty-six new combinations are established
and three are restored. Three species synonymies are restored, one is new, and two are rejected. Across this
diversity of species is a striking diversification of chromosome complements, with X-autosome fusions
occurring at least four times to produce neo-Y sex chromosome systems (X,X,Y and X,X,X,Y), some of
which (Sittisax ranieri and S. saxicola) are sufficiently derived as to no longer preserve the simple traces of
ancestral X material. The correlated distribution of neo-Y and a base autosome number of 28 suggests that

neo-Y origins occurred preferentially in lineages with the presence of an extra pair of autosomes.

Keywords

Amycoida, karyotype, molecular phylogeny, Salticinae, sex chromosomes

Introduction

The jumping spider species long placed in the genus S#tticus Simon, 1901 are well
known in both Eurasia and the Americas as prominent members of habitats as diverse
as boreal forests, marshes, deserts and human habitations (e.g., Locket and Milledge
1951; Prészynski 1968, 1971, 1973, 1980; Harm 1973; Logunov and Marusik 2001).
They belong to the tribe Sitticini, characterized morphologically by the loss of a ret-
romarginal cheliceral tooth, long fourth legs, and an embolus fixed to the tegulum.
Phylogenetic studies have suggested that sitticines arose in the Neotropics, dispersed to
Eurasia, and radiated there (Maddison and Hedin 2003; Maddison 2015), a breadth
of distribution rarely seen in recent lineages of salticids. The Neotropical sitticines
(Figs 1-10) show considerable diversity, with some species having males with colourful
and fringed courtship ornaments (Ad/utticus Galiano, 1987; Figs 2, 3), and others with
shiny metallic colours (Jollas geniculatus group; Figs 7, 113—116). The Eurasian radia-
tion is more sedate in appearance, though there is still diversity in form and markings
in Attulus Simon, 1868 (Figs 15-47).

This work's three goals are to resolve sitticine phylogeny, to review the taxonomy of
sitticines of one region (Canada), and to describe the remarkably diverse chromosomes
of sitticines. Our immediate (and urgent) purpose in studying the group’s phylog-
eny is to settle its turbulent generic classification, which has seen, for instance, some
well-known species change names three times in two years, for example, from Sitticus
floricola (C. L. Koch, 1837) to Sittiflor floricola (by Prészynski 2017a) to Calositticus
floricola (by Blick and Marusik 2018) and back to Sitticus floricola (by Breitling 2019).

Until the last few years, most sitticines were placed in the single widespread
and species-rich genus Sitticus Simon, 1901 (e.g., Platnick 2014 listed 84 species).
Prészynski, who developed our understanding of north-temperate species in a series

Sitticine jumping spiders 3

of papers (1968, 1971, 1973, 1980), recently (2016, 2017a) partitioned this diver-
sity into several genera: Sittipub Prdészynski, 2016, Sittiflor Prészynski, 2017, Sittilong
Prészynski, 2017, Sittisax Proszyniski, 2017, Sittiab Proszynski, 2017, Attulus, and Sit-
ticus. Prészynski did not intend this classification to be phylogenetic, but rather “prag-
matic” (Prészynski 2017b), which is to say, not based on a conceptual justification.
If a classification rejects reference to a broader theory, whether about monophyly or
adaptive zones or predictivity across many characters, then it is not clear what it means,
how it can be tested, or whether it can even be correct, except in its specific statements
about the few characters mentioned. Furthermore, Prészyriski provides little discus-
sion of the diagnostic characters, indeed arguing against explicitly stating or explaining
them (see Prdészynski 2017a; Kropf et al. 2019). Thus, both his characters and his taxa
remain inscrutable.

Breitling (2019) reversed Prészynski’s splitting by synonymizing many of the gen-
era back into Sitticus, based on results from the single mitochondrial gene COI. We
are fortunate that Breitling followed only a small fraction of the implications of his
COI gene tree, for had they been followed more thoroughly they would have yielded
taxonomic chaos in sitticines and throughout salticids, given that they scramble many
well-supported salticid relationships, splitting (for instance) Sitticus sensu lato among
five different tribes (discussed below, with our phylogenetic results). That COI is par-
ticularly bad at resolving salticid phylogeny has been reported previously (Hedin and
Maddison 2001; Maddison et al. 2008, 2014; Bodner and Maddison 2012; Maddison
and Sztits 2019). The results from this single mitochondrial gene therefore have given
us no secure basis for sitticine taxonomy.

Neither Prészynski’s “pragmatic” classification nor Breitling’s COl-based classifica-
tion have promoted taxonomic stability in sitticines. Prdszyniski’s intentionally non-phy-
logenetic approach is particularly problematical. The great majority of systematists no
longer use such “pragmatic” non-evolutionary classifications, as they are not anchored to
a broadly predictive external reality: they are subject to the whims of biologists’ interests
and the character systems they focus on. A taxon delimited for this sense of pragmatism
carries with it no promise of meaning or utility, other than the promise it will bear the
diagnostic characters chosen. Different choices of diagnostic characters would lead to
different classifications, with no basis for selecting among different authors’ approaches
except the weight of authority — in the end, not as pragmatic as a stable phylogenetic
classification, which, by the implications of genetic descent, will predict trait distribu-
tions across the genome. Breitling’s approach might have dampened the instability, as
it is phylogenetic and uses explicit data and analysis, but his choice of the single gene
COI, without supporting morphological information, has yielded a classification in
which we can have little confidence. Prészynski’s and Breitling’s reclassifications might
have been steps forward had they been done in a group of salticids with almost no pre-
vious attention, but the sitticines are reasonably well studied and often mentioned in
the literature. These sudden, comprehensive, conflicting, and largely baseless rearrange-
ments of Sitticus have yielded taxonomic instability in a well-known group.

4 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

Taxonomic instability yields confusion in ecological and other biodiversity litera-
ture about the identity of species studied, and damages the reputation of the taxo-
nomic enterprise. We are now sufficiently capable of resolving phylogeny that we do
not need to rely on the “pragmatic” choices of one authority or on a single misbehaving
gene. Our goal is to provide stronger evidence, explicitly analyzed, for phylogenetic
relationships in order to stabilize the classification of sitticines.

Materials and methods

Morphology

Preserved specimens were examined under both dissecting microscopes and a com-
pound microscope with reflected light. Most of the coquille drawings were done in
1977 or 1978 using a reticle grid in a stereomicroscope. Colour drawings were done in
1974 through 1977 with a stereomicroscope and reticle grid. Pen and pencil drawings
were made recently using a drawing tube on a Nikon MEGOOL compound microscope.
Because some images were made decades ago, we are unable to supply scale bars on
many. Terms used are standard for Araneae. All measurements are given in millimeters.
Carapace length was measured from the base of the anterior median eyes not including
the lenses to the rear margin of the carapace medially; abdomen length to the end of
the anal tubercle. The following abbreviations are used: ALE, anterior lateral eyes; PLE,
posterior lateral eyes; PME, posterior median eyes (the “small eyes”); RTA, retrolateral
tibial apophysis of the male palp.

Specimens were examined from the collections of the American Museum of Natu-
ral History (AMNH), the Canadian National Collection of Insects, Arachnids and
Nematodes (CNC), the Museo Argentino de Ciencias Naturales (MACN), the Muse-
um of Comparative Zoology (MCZ), the Museum of Zoology, Pontificia Universidad
Catdlica, Quito, Ecuador (QCAZ), and the Spencer Entomological Collection of the
Beaty Biodiversity Museum (UBC-SEM).

Nomenclatural authorities

Authors of nomenclatural acts in this paper vary by rank. For acts affecting the syn-
onymy of genera (viz., reinstatement of Attinella and Tomis; synonymies of Sitticus,
Pseudattulus and Sittiab), the authors are those of the paper itself. For all other acts, the
author is W. Maddison. These include the establishment of the Aillutticina, new sub-
tribe, acts that affect the synonymy and placement of species (new synonyms, restored
synonyms, new combinations), and new species.

If not otherwise indicated, the authors of species names are given in the Classifica-
tion section.

Sitticine jumping spiders 5

Molecular phylogeny

Taxa were sampled to cover a diversity of sitticine species groups from Eurasia, North
America, and South America (Table 1). Most were preserved in 95% ethanol, although
we attempted to obtain sequences from some species (Attulus rupicola, A. striatus, A.
cutleri) available only as 70-80% ethanol preserved specimens. We were unable to
obtain sequences from A. striatus and A. cutleri, leaving us with a total of 23 sitticine
species and two outgroups. The outgroups are Breda, from the sister group to sitticines,
and Colonus, from the sister group to remaining amycoids as a whole (see Ruiz and
Maddison 2015; Maddison et al. 2017).

For most samples, DNA was extracted from multiple legs using the Qiagen DNeasy
Blood and Tissue Kit (Qiagen, Valencia, CA) following manufacturer's protocol. Spec-
imens d491 and d492 of Attulus rupicola and d493 of A. zimmermanni were extracted
using standard phenol-chloroform methods. UCE library preparation followed meth-
ods previously used in arachnids (e.g., Starrett et al. 2017; Derkarabetian et al. 2018;
Hedin et al. 2018). Target enrichment was performed using the MYbaits Arachnida
1.1K version 1 kit (Arbor Biosciences; Faircloth 2017) following the Target Enrich-
ment of Illumina Libraries v. 1.5 protocol (http://ultraconserved.org/#protocols).
Libraries were sequenced with an Illumina HiSeq 2500 (Brigham Young University
DNA Sequencing Center) with 150 bp paired end reads. Raw demultiplexed reads
were processed with Puytuce (Faircloth 2016), quality control and adapter removal
was conducted with the ILLUMIpROcEssoR wrapper (Faircloth 2013), and assemblies
were created with VELvET (Zerbino et al. 2008) at default settings. The Szttilong lon-
gipes ARV4504 sample was sequenced on a NovaSeq 6000 at the Bauer Core Facility
at Harvard University with 150 bp paired end reads, and was assembled with TRINITY
(Grabherr et al. 2011) with default settings. Contigs were matched to probes using
minimum coverage and minimum identity values of 80. UCE loci were aligned with
MAFFT (Katoh and Standley 2013) and trimmed with Gstocxs (Castresana 2000;
Talavera and Castresana 2007), using -- bl 0.5, --b2 0.5, --b3 10, --b4 4 settings in
the PHyLuce pipeline.

In the resulting set of loci, most taxa have over 100,000 base pairs of sequence
data, but some are less thoroughly sequenced. ‘The less thoroughly sequenced taxa are:
J. leucoproctus d478 (13,943 bp), Attulus rupicola d491 (46,660 bp), Attulus rupicola
d492 (65,500 bp), and A. zimmermanni d493 (68,285 bp). The last species is repre-
sented by an alternative well-sequenced specimen, the others by well-sequenced close
relatives. Although we did analyses with the entire set of taxa (“All Taxa’), we were
concerned that the weakly sequenced taxa would disrupt resolution. Therefore, we
rely primarily on analyses (and bootstrap values) that exclude these and use only the
remaining well-sequenced taxa (“Core Taxa’). The Core Taxa dataset also excludes the
less thoroughly sequenced of the two specimens of Jollas cupreus (d473, 92,549 bp).

This pipeline therefore resulted in two collections of genes, one of 968 loci for
all the taxa (“All Taxa”), the other of 957 loci for the core set of well-sequenced taxa

6 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

Table |. Specimens from which UCE sequence data gathered. “UCE loci” indicates number of loci from
Puytuce. “Reads Pass QC” indicates number of reads retained after quality control and adapter removal
via Illumiprocessor.

Species Specimen sex Locality Reads Pass QC Contigs UCE loci
Aillutticus nitens d475 f Uruguay: Canelones: -34.867, -56.009 946351 207743 434
Attinella dorsata d490 m U.S.A.: California: 37.2834, -120.8515 1617332 360661 480
Attulus ammophilus d482 m Canada: British Columbia: 49.7963, -119.5338 1471891 351670 588
A. burjaticus RU18-7302f Russia: Tuva: 50.205, 95.135 529905 151897 627
A. distinguendus RU18-6432 f Russia: Tuva: 50.746, 93.142 406186 90846 626
A, fasciger d487 m Canada: Ontario: 43.35074, -79.75928 1370738 299273 564
A, finschi d480 m Canada: British Columbia: 49.0261, -114.0611 1489551 303924 579
A, floricola d488 m Canada: Saskatchewan: 52.4898, -107.3843 1466702 303612 606
A, inexpectus RU18-6799 m Russia: Tuva: 50.669, 92.9844 261947 60612 653
A, longipes ARV4504. sm Italy: Stilfs 16385503 42677 515
A. mirandus RU18-7308 f Russia: Tuva: 50.205, 95.135 468358 110900 649
A. pubescens d483 m Canada: British Columbia: 49.2, -123.2 1316697 279173 503
A, rupicola d491 m Poland: Cisna near Lesko 187507 58418 312
A, rupicola d492 m Poland: Bukowksa Kopa 418777 137114 397
A. saltator d512 m Germany: Saxony: 51.607, 12.711 416618 113416 591
A, sylvestris d489 m U.S.A.: California: 36.3646, -121.5544 1289981 278727 506
A. terebratus RU18-5346 m Russia: Novosibirsk Oblast: 53.73, 77.866 306744 72547 668
A. zimmermanni d493 m Poland: Grabarka 52.417, 23.005 338718 113167 408
A. zimmermanni RU18-5156 m Russia: Novosibirsk Oblast: 53.721, 77.726 435654 93640 627
Breda bicruciata d471 f Uruguay: Lavalleja: -34.426, -55.195 646088 248616 549
Colonus hesperus d472 m ULS.A.: Arizona: 34.5847, -112.5707 1015130 250378 448
Jollas cellulanus d479 f Argentina: Neuquén: -37.0679, -69.7566 981935 268639 497
J. cupreus d473 m Ecuador: Orellana: -0.526, -77.418 1419103 289905 469
J. cupreus d474 m Ecuador: Orellana: -0.526, -77.418 3513351 723782 607
J. leucoproctus d478 f Uruguay: Maldonado: -34.94, -54.95 121131 61298 109
Sittisax ranieri d481 m U.S.A.: Oregon: 44.0322, -121.6722 1529835 322636 536
Tomis manabita d476 m Ecuador: Manabi: -1.5497, -80.8104 2524270 710859 651
T. palpalis d477 m Ecuador: Napo: -0.1996, -77.7023 1211674 256367 582

(“Core Taxa’). A filter of occupancy was then applied, eliminating all loci which had
sequences for fewer than seven of the 20 well-sequenced taxa of the ingroup (Jollas,
Attinella, Tomis, Sittisax, Attulus), resulting in 810 loci in the All Taxa dataset and
803 in the Core Taxa dataset. Preliminary analyses of these loci revealed some whose
gene trees strongly suggested two paralogs or chimeras were included: a single very
long branch isolating a few taxa (which for all other considerations and subsequent
analyses showed no indication of being so distinctive or related to one another), whose
sequences differed from the others extensively and consistently. Out of caution we
chose to discard a locus if its preliminary gene tree (RAxML 8.2.8, Stamatakis 2014,
single search, default settings) had the longest branch at least five times longer than the
second longest branch. Inspection of the results indicated this matched approximately
our subjective judgment of a strong suspicion of paralogy. This filter left 749 loci in the
All Taxa dataset and 757 loci in the Core Taxa dataset.

Maximum likelihood phylogenetic analyses were run using IQ-TREE version
1.6.7.1 (Nguyen et al. 2015), run via the Zephyr package (version 2.11, Maddison
and Maddison 2018a) of Mesquite (Maddison and Maddison 2018b). The data were
analyzed both without partitions (“unpartitioned”) and partitioned by locus, allow-

Sitticine jumping spiders 7

ing the possibility of separate rates and substitution models (Kalyaanamoorthy et al.
2017). We ran 50 separate search replicates for the maximum likelihood tree for the
concatenated analysis. We performed a standard bootstrap analysis with 1000 repli-
cates and the same model and partition settings.

A separate small phylogenetic analysis was done to explore the distinction in At-
tulus floricola between hemispheres, using data of other specimens in Genbank and
BOLD (boldsystems.org), to blend with our data. Insofar as only COI barcode data
are available online, and this gene struggles to reconstruct salticid phylogeny (Hedin
and Maddison 2001; Maddison et al. 2008, 2014; Bodner and Maddison 2012; Brei-
tling 2019; Maddison and Sztits 2019), we provided a skeletal constraint tree of our
UCE specimens from which we could obtain COI data, so that the gene’s burden
would be only to place the extra COI-only floricola group specimens on this skeleton.
We obtained COI data for our UCE taxa by mining the UCE reads for COI-alignable
bycatch. A local database was assembled in Geneious v1 1.0.4 comprising labeled VEL-
vet UCE contigs for all sequenced taxa, then published A. striatus sequences (voucher
BIOUG14302-A06) were used to query this local database using BLASTN (max e
value of 1x10°'°). Retaining only high-coverage sequences, we recovered COI bycatch
for all taxa except for A. saltator, A. inexpectus, and A. rupicola. For A. saltator and A.
rupicola we substituted COI data from Genbank from another geographically proxi-
mate specimen. The constraint tree was set to match Figure 48 in topology. Then, we
added and aligned COI sequences of A. floricola from scattered locations, as well as
specimens of A. caricis (from the Netherlands) and A. sylvestris (from Canada). (The
latter were identified in BOLD as A. rupicola, but inspection of genitalic photographs
courtesy of G. Blagoev shows they are A. sylvestris.) The gene tree was recontructed
by RAxML (Stamatakis 2014), with codon positions as separate partitions, and using
Figure 48 as a skeletal (partial) constraint tree.

Sequence reads are deposited in the Sequence Read Archive (BioProject submis-
sion ID PRJNAG605426, http://www.ncbi.nlm.nih.gov/bioproject/605426). Align-
ments and trees are deposited in the Dryad data repository (https://doi.org/10.5061/
dryad.cjsxksn2q).

Chromosomes

Chromosomes were studied in 17 taxa of Sitticina. The specific identity of the speci-
men labelled “A. rupicolalfloricola’ is ambiguous because the voucher specimen has not
been located, and the first author is not confident he was able to distinguish the two
species in the 1980s. Although its specific identity is not known, it can be confidently
placed within the floricola group, and so can play a role in phylogenetic interpretation.

Meiotic chromosomes were observed in testes of adult and subadult males using
Feulgen staining, following the methods of Maddison (1982), except that no colchi-
cine was used. Most preparations of Nearctic material were done between 1980 and
1989, and scored for autosome number and form and sex chromosome system soon
thereafter. In the years since, some of the slides have faded considerably, and even

8 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

with phase contrast they can no longer be scored. For most species we were able to
confirm the old scores through re-examination (in an Olympus BX51 phase contrast
microscope), except as noted in Chromosome observations. Because of the long his-
tory of this study, our photographs are of varied ages and qualities. We recognize that
chromosome scoring of some species has uncertainty, and that future studies should
be directed to confirming or correcting our intepretations. Nonetheless, the broad pat-
terns we describe are supported even taking the uncertainty into account.

Evidence for scoring chromosome complement of each species is described in
Chromosome observations. Most chromosome scoring was done from meiotic nu-
clei in first metaphase or diakinesis showing chromosomes that are well separated, or,
if overlapping, easily interpretable. Although well-spread mitotic nuclei would have
added useful data, we judge meiotic chromosomes to be sufficient as they show distinc-
tive features, e.g. when they are oriented by the centromere pulling toward the pole on
the metaphase plate. Metacentrics show an obvious bend at the centromere where the
second arm hangs loose like a dog’s ear (Fig. 130), while acrocentrics show an opposite
bend more distally (at chiasmata), or no bend (if chiasmata are terminal), and a nar-
rower neck to the centromere stretched pole-ward (Figs 131, 140, 143, 147, 154, 156,
164). In most specimens, multiple nuclei contributed to the scoring. In other salticids
(e.g., Maddison 1982), the Xs of the X,X,0 sex chromosome system have distinctive
behaviour during meiosis. At first metaphase they typically lie toward one pole, side
by side and without chiasmata. They are heteropycnotic, condensing early, but by first
metaphase slightly decondensed, and in second prophase condensed. We use this be-
haviour as evidence for interpreting chromosomes as Xs, or for interpreting portions
of chromosomes as representing ancestral X material. For several species additional
evidence came from metaphase II counts, and for one (Sittisax ranieri) female mitosis
in subadult digestive glands was examined.

In describing chromosome complements, we use “a” and “m” to indicate one-
armed (acrocentric/telocentric) and two-armed (metacentric/submetacentric) chro-
mosomes respectively. Thus, “26a+XaXa0” would mean “26 acrocentric autosomes
plus two X’s, both of which are acrocentric”. In all cases, the multiple Xs of a male are
interpreted as not being homologous, and therefore it would be more proper to refer

to the systems as XO) A 2, Or X,X,X,Y rather than as XX0, XXY, or XXXY. How-

D> 6

ever, the “1”, “2”, “3” will be left implicit, omitted for ease of reading, to avoid overly

complex labels like Xa,Xa,Xa,Ym.

Phylogenetic results

The maximum likelihood tree from the UCE data is shown in Figure 48, which incor-
porates results from both partitioned and unpartitioned analyses. As seen in previous
results from fewer genes (Ruiz and Maddison 2015), Aillutticus Galiano, 1987 is the
sister group to all other sitticines sampled. Ai//utticus is the only sampled representa-
tive of what is likely a large radiation of little-studied Neotropical sitticines with high,
rounded carapaces and unusual genitalia, currently including five genera (Galiano

Sitticine jumping spiders 9

1987; Ruiz and Brescovit 2005, 2006). As described under classification, we propose
the name Aillutticina, new subtribe, for the Ai//utticus group of genera, and the name
Sitticina for the remaining sitticines.

The phylogeny of Sitticina shows two major groups, the Jollas- Tomis clade and Attulus.
The Jollas-Tomis clade is distributed entirely in the Americas except for the two species
of Sittisax, Attulus is entirely Eurasian except for 8 species in North America. The only
previously published comprehensive phylogeny of sitticines, of Prdészynski (1983), is sub-
stantially similar in placing Sittisax and A‘tinella outside of the major clade of the floricola,
distinguendus and penicillatus species groups. The most notable differences between his ar-
rangement and ours are the placements of Attulus pubescens and.A. dzieduszycki. Prészynski’s
more recent (2017a) classification into genera, however, is discordant in many respects with
our results, as can be seen in the many combinations that we establish or reinstate below in
order to achieve monophyly of genera and subgenera. This discord may have arisen partly
because Prdészynski was not attempting to create a taxonomy that reflected phylogenetic
relationships, but rather the distribution of a few diagnostic characters (Prészynski 2017b).

Our UCE phylogeny differs in several respects from Breitling’s (2019) COI phylog-
eny. Ours places Sittisax ranieri next to Tomis, distant from the Eurasian Radiation, while
his places it next to Attulus finschi. The other disagreements are not visible in the isolated
portion of the tree shown in Breitling’s figure 9B, but are visible in his more complete
supplemental figure “Salticidae”. It places Attinella concolor sister to the euophryine Si-
dusa, Attulus fasciger among the plexippines, Tomis manabita (“Sitticus sp. MCH-2003”)
as sister to the asemoneine Asemonea, and Jollas cupreus as sister to the lapsiine Thrandina
— thus mixing the sitticines among three different subfamilies and 5 tribes. Given our
far stronger data (hundreds of loci, multiple linkage groups, many times more nucleo-
tide sites), inclusion of more Neotropical sitticines, more efficient analysis (likelihood as
opposed to neighbour joining), and concordance with morphological traits uniting the
sitticines, we consider Breitling’s phylogeny to be in error. The startling scrambling of
established clades in Breitling’s supplemental figures is in accord with previous studies in
salticids, which have shown the COI gene to be particularly error-prone in reconstruct-
ing phylogeny (Hedin and Maddison 2001; Maddison et al. 2008, 2014; Bodner and
Maddison 2012; Maddison and Sztits 2019). However, our phylogeny agrees with one
important result from Breitling’s study: the close relationship of A. pubescens with A. ter-
ebratus (though, as noted above, their close relative A. fasciger is placed in another tribe).

Although Aztulus includes some Nearctic members, it is considerably more species-
rich in Eurasia, and is most parsimoniously interpreted as having radiated there. The
few Nearctic members of this clade are likely recent returns from the Palearctic, insofar
as they are Holarctic (Attulus floricola, A. cutleri, A. finschi), close relatives of Eurasian
species (A. sylvestris within the A. floricola group, A. striatus close to A. rivalis), or recent
introductions (A. ammophilus, A. fasciger, A. pubescens: see Proszynski 1976, 1983 and
Cutler 1990). Our results thus support Prdészynski’s (1983) hypothesis of a Palearctic
radiation of Sitticus sensu lato, although we differ in concluding that only one sub-
group diversified in Eurasia, Attulus, arising from an earlier Neotropical diversification.

The deep branches of the Eurasian Radiation are short, suggesting the group di-
versified rapidly. Nonetheless, the monophyly of subgenus Sitticus is well supported

10 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

by a bootstrap percentage of 100 in our primary Core Taxa analysis (Fig. 48). The
monophyly of subgenus Aztu/us has weak bootstrap support in the partitioned analysis
(72%), although good support in the unpartitioned analysis (95%). As well, the major
subgroup of subgenus Aztulus excluding A. saltator and A. mirandus is well supported
(92% or 96%). Despite its weak support in the partitioned analysis, monophyly of
subgenus AZtulus as a whole is consistent across multiple analyses, for example, when
the filter for loci present in at least seven core taxa is changed from seven to four or
ten. Analyses (following the same methods described above) without A. longipes gave
99.6% bootstrap support to subgenus Aztulus.

The relationships among Aztulus species are concordant with morphological ex-
pectations with one exception: the placement of A. burjaticus with A. zimmermanni,
suggesting that the longer embolus of A. zimmermanni and the floricola group are
convergent. Otherwise, the floricola group holds together, as do the morphologically
similar pairs of A. ammophilus/distinguendus and mirandus/saltator. The placement of
A, pubescens nested within the terebratus group indicates that the very short embolus of
the former is a derivation from the very long embolus of the latter.

Jollas and Tomis together form a Neotropical radiation and share (typically) an
RTA that appears displaced basally, so as to appear to arise closer to the patella, as well
as anteriorly placed epigynal openings.

Classification

The phylogenetic results lead us to revise the generic division of sitticines. Unless
we are to put all Sitticina into a single genus, perhaps palatable for the shallow-
diverging Eurasian fauna, but not for the deep Neotropical lineages, then Tomis
must be restored for many of the Neotropical species. Given that, Sittisax must be
separated from Attulus/Sitticus, rejecting Breitling’s synonymy of this taxon with
Sitticus. These choices are relatively easy. The more difficult choices concern the
Eurasian Radiation.

Here we give a taxonomic review of the tribe, focussing especially on the species
in Canada, and the two new species used in the molecular phylogeny (Jollas cupreus
and Jomis manabita). In order to faciliate the use of figures for identification and com-
parison of species in North America, the sequence of taxa in figures will be different
from that in the text, with a series of standardized plates placing images of all of the

Canadian species in a block (Figs 49-103).

Tribe Sitticini Simon, 1901

Amycoid salticids with fourth legs much longer than third and retromarginal cheliceral
tooth lacking. Ancestrally they were ground-dwellers in the Neotropics, later diversi-
fying in Eurasia to include species that live on tree trunks (e.g., A. finschi) and up in
vegetation (e.g., Aztulus floricola).

Sitticine jumping spiders 11

Eleven genera are here recognized in the Sitticini, including one (Semiopyla Simon,
1901) whose placement is unclear, and thus remains incertae sedis within the tribe. Two
genera are in Eurasia (Aztulus and Sittisax), while a disjunct set of eight genera are in
South America (the five aillutticines, plus Toms, Jollas, and Semiopyla). This geographi-
cal partitioning matches a phylogenetic division approximately, but not precisely, for
the Holarctic Sittisax is phylogenetically a member of the Neotropical radiation. North
America has four genera, one arising from the Eurasian radiation (Attulus), and three
from the Neotropical radiation (Attinella, Sittisax, and Tomis).

Despite the synonymy of Sitticus with Attulus, the names Sitticini and Sitticina can

persist (ICZN Article 40.1).

Subtribe Aillutticina W. Maddison, new subtribe
http://zoobank.org/4DBE8F82-300A-4AE0-9A1 1-7A0DC55D7099

Figures 1-4

Type genus. Ai//utticus Galiano, 1987

Diagnosis. This group of five Neotropical genera was first recognized by Ruiz and
Brescovit (2005, 2006), who characterize it as sharing “a high, broad carapace, laterally
rounded behind the posterior lateral eyes, and the slightly convex dorsal surface of the
cephalic region”. The contained genera are:

Aillutticus Galiano, 1987
Amatorculus Ruiz & Brescovit, 2005
Capeta Ruiz & Brescovit, 2005
Gavarilla Ruiz & Brescovit, 2006
Nosferattus Ruiz & Brescovit, 2005

Subtribe Sitticina Simon, 1901

There are no known morphological synapomorphies of this subtribe, but the molecu-
lar data show clearly that the five genera listed here form a clade. ‘There are two major
subgroups according to the UCE phylogeny: the genus Aztulus, a primarily Eurasian
radiation, and the Jollas- Tomis clade (Attinella, Jollas, Sittisax, Tomis), a primarily Neo-
tropical radiation. We divide the taxonomy below into those two major groups, and
under each discuss the genera, describe the Canadian species and two new Ecuadorian
species used in the molecular work.

Genus Aztulus Simon, 1868, restored (to respect its priority over Sitticus)

Attulus Simon, 1868 (type species Attus helveolus Simon, 1871)
Sitticus Simon, 1901 (type species Avaneus terebratus Clerck, 1757)

12 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

Figures I-14. Subtribe Aillutticina (1-4) and the /ollas-Tomis clade of the subtribe Sitticina (5-14)
1-4 Aillutticus nitens, Uruguay (-34.877, -56.023): 1-3 male 4 female 5,6 Yomis palpalis male and fe-
male, Ecuador (-0.1996, -77.7023) 7, 8 Jollas species: 7 J. cupreus male, Ecuador (-0.675, -76.397) 8 Jol-
las sp. female, Ecuador (-0.7223, -77.6408) 9 J. leucoproctus, Uruguay (-34.94, -54.95) 10 /. flabellatus,
Uruguay (-34.426, -55.195) Ll=-14 AZtinella dorsata male (11-13) and female (14), Canada (48.870,
-123.379). Also included in the Jollas-Tomis clade is Sittisax (Figs 99-103). Additional members of the
Jollas- Tomis clade can be seen in Figs 108-128.

i a ihenc

Sitticulus F. Dahl, 1926 (type species Aitus saltator O. Pickard-Cambridge, 1868), syn. nov.
Calositticus Lohmander, 1944 (type species Attus caricis Westring, 1861), syn. nov.
Hypositticus Lohmander, 1944 (type species Aranea pubescens Fabricius, 1775), syn. nov.
Sittipub Prészynski, 2016 (type species Aranea pubescens Fabricius, 1775), syn. nov.
Sittiflor Prészynski, 2017 (type species Euophrys floricola C.L. Koch, 1837), syn. nov.
Sittilong Proszynski, 2017 (type species Attus longipes Canestrini, 1873), syn. nov.

Sitticine jumping spiders 13

We unite the primary Eurasian radiation under the single genus Attulus because of
the recency of the radiation, the very short phylogenetic branches separating the sub-
groups, and the clade’s morphological homogeneity. The total phylogenetic depth of
Attulus is far less than that of its sister group (Fig. 48), but more importantly, the deep-
est branches of Attulus are very short. This suggests a rapid radiation, and that any sub-
groups will have only limited predictive value about traits, as most of the divergence
occurred since the initial radiation. The monophyly of the major subgroups is to some
extent uncertain, and so any generic division could be unstable. The morphological
diversity encompassed by Aftulus (e.g. variation in narrowness of carapace, leg length,
embolus length, position of epigynal openings) is arguably less than that of other sta-
ble genera like Pellenes and Habronattus; the subgenera we recognize are comparable
to species groups in Habronattus (Maddison and Hedin 2003) or subgenera in Pellenes
(Logunov et al. 1999). By considering Aztulus as a single genus with subgenera, we also
simplify identifications by ecologists and others. A Eurasian salticid, even a juvenile,
can easily be keyed to Attulus based on the long fourth legs and absence of retromar-
ginal cheliceral teeth, except only for the exclusion of Sittisax.

Our choice to consider all but two Eurasian species as belonging to Attulus is in-
formed partly by their phylogenetic context among Neotropical salticids. From a
Palearctic perspective, the Eurasian radiation of sitticines may seem to represent a line-
age of salticids so distinctive and species-rich that they deserve splitting into many gen-
era, especially since the sister group of sitticines among the Old World salticids is the
huge clade Salticoida (Maddison 2015), which is divided into hundreds of genera. From
the Americas, though, the Eurasian sitticine radiation appears as a shallow expatriate
lineage, the tip of the iceberg of a large and deeply diverging Neotropical radiation (the
Sitticini, and more broadly, the Amycoida). If more generic subdivision is needed, it will
be in the much more divergent and poorly explored sitticine fauna of South America.

The appropriate name for this unified genus is Attulus, as it is far older than Sitzi-
cus, and has been used continuously, though for only a few species. Two proposals have
been made to ignore priority and instead use Sitticus, the generic name used for most of
the species until Prészyrski’s (2016, 2017) splitting. Prészynski himself had proposed
to the ICZN in 2008 suppression of Attulus in favour of Sitticus, but in 2018 appar-
ently withdrew that proposal (ICZN 2018). Breitling (2019) also proposed that the
younger name Sitticus be used. We argue that priority in general should be respected
unless it would disrupt a long-stable name against a little-used alternative. In this case,
Sitticus has already been destabilized, Aztu/us has been used more or less continuously,
and most species have already been moved to Attulus by Prészynski. The World Spider
Catalog (WSC 2019) and other resources (Metzner 2019) have already begun to use
Attulus for most species. Abandoning nomenclatural rules to avoid facing the conse-
quences of new information will over the long term likely lead to instability or to clas-
sifications based on the weight of authority, just as with abandoning monophyly. Thus,
the least disruptive choice is to use the name “Aztulus”.

However, there is value in offering a weaker recognition of three subgroups of Az
tulus, as subgenera, given that there are names available: Aztulus, Sitticus, and Sittilong.
Our results support reciprocal monophyly of the subgenera Attulus and Sitticus, and

14 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

a placement of Sittilong outside of both. Monophyly of subgenus Aztulus has variable
bootstrap support (72% to 95%, Fig. 48), although the clade’s presence is consistent
across various alternative analyses (when Sittilong is not included; when the filter for
loci present in at least seven core taxa is changed to four or ten). Even if subgenus At
tulus falls apart with more data, the bulk of the subgenus would likely hold together, as
there is high bootstrap support for the large subclade including the type species A. dis-
tinguendus. The low bootstrap support for the subgenus as a whole (in the partitioned
analysis) derives from the weakness of inclusion of the unusual penicillatus group (rep-
resented by A. saltator and A. mirandus; see Logunov 1993), which might eventually
need a separate subgenus (for which a name, Sitticulus F. Dahl 1926, is available).

The three subgenera have subtle but mostly consistent morphological differences.
Attulus s. str. tends to have smaller and more compact bodies, with roundish carapaces
(Figs 15-38). Sitticus have a narrower carapace and longer legs (Figs 39-47), and (ex-
cept in A. relictarius) a large sweeping retrolateral tibial apophysis (Figs 74, 79, 84).
Sittilong is notable for its long first legs.

Attulus includes 49 species in three subgenera:

Subgenus Aztu/us Simon, 1868, with 41 species:
Attulus (Attulus) albolineatus (Kulczyhski, 1895), comb. nov., transferred from Sitticus
Attulus (Attulus) ammophilus (Thorell, 1875)
Attulus (Attulus) ansobicus (Andreeva, 1976)
Attulus (Attulus) atricapillus (Simon, 1882), comb. nov., transferred from Calositticus
Attulus (Attulus) avocator (O. Pickard-Cambridge, 1885)
Attulus (Attulus) barsakelmes (Logunov & Rakov, 1998), comb. nov., transferred
from Sitticus
Attulus (Attulus) burjaticus (Danilov & Logunov, 1994)
Attulus (Attulus) caricis (Westring, 1861), comb. nov., transferred from Calositticus
Attulus (Attulus) clavator (Schenkel, 1936)
Attulus (Attulus) cutleri (Prészyfski, 1980), comb. nov., transferred from Calositticus
Attulus (Attulus) damini (Chyzer, 1891)
Attulus (Attulus) distinguendus (Simon, 1868) (= type species Attus helveolus Simon, 1871)
Attulus (Attulus) dubatolovi (Logunov & Rakov, 1998)
Attulus (Attulus) dudkoi (Logunov, 1998), comb. nov., transferred from Calositticus
Attulus (Attulus) dzieduszyckii (L. Koch, 1870), comb. nov., transferred from Sittisax
Attulus (Attulus) eskovi (Logunov & Wesolowska, 1995), comb. nov., transferred
from Sitticus
Attulus (Attulus) floricola (C. L. Koch, 1837), comb. nov., transferred from Calositticus
Attulus (Attulus) goricus (Ovtsharenko, 1978)
Attulus (Attulus) hirokii Ono & Ogata, 2018
Attulus (Attulus) inexpectus (Logunov & Kronestedt, 1997), comb. nov., transferred
from Calositticus
Attulus (Attulus) inopinabilis (Logunov, 1992)
Attulus (Attulus) karakumensis (Logunov, 1992)

Sitticine jumping spiders 15

Attulus (Attulus) kazakhstanicus (Logunov, 1992)
Attulus (Attulus) mirandus (Logunov, 1993)
Attulus (Attulus) monstrabilis (Logunov, 1992), comb. nov., transferred from Calositticus
Attulus (Attulus) nenilini (Logunov & Wesolowska, 1993)
Attulus (Attulus) nitidus Hu, 2001, comb. nov., transferred from Sitticus
Attulus (Attulus) niveosignatus (Simon, 1880)
Attulus (Attulus) penicillatus (Simon, 1875)
Attulus (Attulus) penicilloides (Wesolowska, 1981)
Attulus (Attulus) pulchellus (Logunov, 1992), comb. nov., transferred from Calositticus
Attulus (Attulus) rivalis (Simon, 1937), comb. nov., and removed from synonymy
with A. striatus (Emerton).
Attulus (Attulus) rupicola (C. L. Koch, 1837), comb. nov., transferred from Calositticus
Attulus (Attulus) saltator (O. Pickard-Cambridge, 1868)
Attulus (Attulus) sinensis (Schenkel, 1963)
Attulus (Attulus) striatus (Emerton, 1911), comb. nov., transferred from Calositticus
Attulus (Attulus) sylvestris (Emerton, 1891), comb. nov., transferred from Sitticus,
removed from synonymy with A. palustris
Attulus (Attulus) talgarensis (Logunov & Wesolowska, 1993)
Attulus (Attulus) vilis (Kulczytiski, 1895)
Attulus (Attulus) zaisanicus (Logunov, 1998)
Attulus (Attulus) zimmermanni (Simon, 1877), comb. nov., transferred from Calositticus
Subgenus Sitticus Simon, 1901, with seven species:
Attulus (Sitticus) fasciger (Simon, 1880), comb. nov., transferred from Sitticus
Attulus (Sitticus) finschi (L. Koch, 1879), comb. nov., transferred from Sitticus
Attulus (Sitticus) godlewskii (Kulczyniski, 1895), comb. nov., transferred from Sitticus
Attulus (Sitticus) pubescens (Fabricius, 1775), comb. nov., transferred from Sitticus
Attulus (Sitticus) relictarius (Logunov, 1998), comb. nov., transferred from Sitticus
Attulus (Sitticus) tannuolana (Logunov, 1991), comb. nov., transferred from Sitticus
Attulus (Sitticus) terebratus (Clerck, 1757) (type species of Sitticus), comb. nov.,
transferred from Sitticus
Subgenus Sittilong Prészynski, 2017, with one species:
Attulus (Sittilong) longipes (Canestrini, 1873) (type species of Sittilong), comb. nov.,

transferred from Sittilong

Subgenus Aztulus Simon, 1868
Figures 15-38, 49-73

Attulus Simon, 1868 (type species Attus helveolus Simon, 1871).

Sitticulus F. Dahl, 1926 (type species Attus saltator O. Pickard-Cambridge, 1868).
Calositticus Lohmander, 1944 (type species Attus caricis Westring, 1861).

Sittiflor Prészynski, 2017 (type species Euophrys floricola C.L. Koch, 1837).

16 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

Body generally more compact than in subgenus Sitticus, with a wider carapace. The
spermatheca is a simple tube, folded near the middle. From the point at which the
copulatory ducts enter the spermatheca, the spermatheca extends medially to the ferti-
lization duct, but also laterally and then posteriorly (floricola group) or medially (most
others) to a separate posterior lobe. Most Attulus (Attulus) have the embolus short,
arising near the basal prolateral corner of the bulb, and the tegulum with basal edge
more or less straight (not rounded). Several species have a rounder bulb and longer em-
bolus, representing two or three lineages: the floricola group (A. caricis, A. floricola, A.
inexpectus, A. rupicola, A. sylvestris), the striatus group (A. striatus, A. rivalis, A. cutleri,
A, dudkoi) and the zimmermanni group (A. zimmermanni, A. atricapillus). These also
have the folded spermathecae rotated slightly compared to the other Attulus, with the
posterior lobe pointing posteriorly, rather than medially. The placement of A. niveosig-
natus in Attulus (Attulus) is somewhat doubtful, as the position of the tibial apophysis
and the anterior medial epigynal openings both resemble those of Sittisax and Attulus
subgenus Sittilong. We are reluctant to move it, however, until it is better studied.

Five species of Attulus (Attulus) are known from North America, all of which occur
in Canada, as follows.

Attulus (Attulus) ammophilus (Thorell, 1875)
Figures 27—30, 69-73

Attus ammophilus Thorell, 1875

Remarks. Attulus ammophilus is part of the species-rich distinguendus group that is
otherwise unrepresented in North America. We have collected it from rocks on the
ground in Ontario, British Columbia, and Utah, on litter among marsh plants along
the edge of a lake in Siberia, and occasionally from buildings. It was introduced into
North America during the 20" century (Prészyriski 1976, 1983).

Material examined (all in UBC-SEM): Canapa: Ontario: Hamilton (69 males,
35 females), Oakville (4 males, 3 females), Toronto (1 male), Windsor (1 male, 2 fe-
males); British COLUMBIA: 49.7963, -119.5338 (1 male, 2 females), 49.95, -119.401
(3 males, 2 females); U.S.A.: Urau: 40.7482, -112.1856 (5 males, 7 females), 40.7672,
-112.1575 (2 males).

Attulus (Attulus) floricola (C.L. Koch, 1837)
Figures 33-35, 49-53

Euophrys floricola C. L. Koch, 1837.

Attus palustris Peckham & Peckham, 1883 (specimens in MCZ labelled as types, ex-
amined, but see below).

Attus morosus Banks, 1895 (synonymized by Prdészyfiski 1980; confirmed here by ex-
amination of holotype female in MCZ from Olympia, Washington).

Sitticine jumping spiders 17

ibe, 12

r Attulus
ammaphilus

93.142) 18-20 male and female A. mirandus, Tuva (50.205, 95.135) 21-23 A. burjaticus: 21 male,
Tuva (50.68, 92.99) 22 male, Tuva (50.205, 95.135) 23 female, Tuva (50.68, 92.99) 24={26 A. zim-
mermanni: 24, 25 male Novosibirsk Oblast (53.721, 77.726) 26 female Novosibirsk Oblast (53.730,
77.865) 27-30 A. ammophilus: 27 male Tuva (50.6690, 92.9844) 28 male Ontario, Oakville 29 female
Ontario, Hamilton 30 male British Columbia (49.08, -119.52). For additional images of A. ammophilus,
see Figs 69-73. For additional images of Attulus (Attulus), see Figs 31-38, 49-73.

Remarks. A widespread Holarctic species often found in retreats in dry flower heads in
wetter areas such as marshes, A. floricola is distinctive for the sharp white lines around
the eyes of males, forming an apparent mask (Fig. 34). Attulus floricola has often been
confused in the past with its close relatives, but the distinctions have been clarified
considerably by Prészynski (1980) and Logunov and Kronestedt (1997).

18 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

We treat the North American populations as full floricola, not a distinct subspe-
cies. While Nearctic populations were long recognized as a separate species palustris,
Prészyniski (1980) suggested they are conspecific with the Eurasian populations. He
maintained them as a distinct subspecies, but he expressed doubt as to whether even
that distinction was warranted. We concur with his skepticism. If any consistent differ-
ences exist between the continents, they are no more visible than any differences that
might exist between the Eurasian and North American populations of other species
for which we don’t recognize subspecies such as Sittisax ranieri, Attulus cutleri, Den-
dryphantes nigromaculatus (Keyserling, 1885), Pellenes ignifrons (Grube, 1861), and
Pellenes lapponicus (Sundevall, 1833).

The results of our COI analysis of Palearctic and Nearctic floricola group (Fig. 104)
show all floricola to be close on the gene tree, with the New World specimens in two
clades (not clearly related to one another) and the German specimens in a third clade.
This suggests that A. floricola is not cleanly or deeply divided between the Nearctic and
Palearctic. The molecular and morphological evidence leads us to fully synonymize
palustris into floricola.

Within North America, the characterization of A. floricola has been muddied by
confusion with a second species, A. sylvestris. Attulus sylvestris, long synonymized with
palustris, is a distinctively different species. Attulus floricola is larger-bodied, has a much
more contrasting colour pattern, and longer legs. Attulus floricola has a different angle
of the spermaphore loop (subtle but consistent; Fig. 49 vs. Fig. 54), and in females
the darkness of the spermathecal lobe is visible through the anteriormost portion of
the epigynal atrium (Fig. 50 vs. Fig. 55). Attulus sylvestris has genitalia more similar
to those of the Eurasian A. caricis, A. rupicola, and A. inexpectus, as noted below. The
synonymy of sylvestris with palustris was originally proposed by Peckham and Peckham
(1909), after which Kaston (1948) may have stirred confusion by choosing to illustrate
palustris using Emerton’s (1891) figure of sylvestris.

A more serious confusion apparently occurred with the labelling of type specimens
of Attus palustris. The description by Peckham and Peckham (1883) refers without
doubt to the common white-striped species long known as Sitticus palustris (Fig. 34):
males dark brown, reddish toward eyes, marked with white lines, including those
around the eyes, and palp with some white hairs on several segments of the palp. As
well, the habitat suggested by the name “palustris” is marsh or swamp, more typical
for A. floricola than A. sylvestris. However, the specimens labelled as the types of Aztus
palustris in the MCZ are clearly specimens of the less common dusty brown species
(i.e., Emerton’s sylvestris, Fig. 32). These specimens, we argue, are mislabelled: they do
not match the Peckhams’ description, and thus are not the type specimens of A. palus-
tris. That the Peckhams viewed the white-striped form as typical palustris can be judged
not only from their 1883 description, but also from their implicitly distinguishing two
forms in their 1909 statement “Mr. Emerton agrees with us that the form which he
described as sylvestris is a variety of palustris, with the leg a little shorter and stouter.”
The label of the holotype does not appear to be in the handwriting of either George
or Elizabeth Peckham, and it is possible that these “types” were so labelled after 1883.

Sitticine jumping spiders 19

a4 -
Attulus sylvestris

Figures 31-38. Azzulus subgenus Aztulus, continued (floricola group) 31, 32 Aitulus sylvestris: 31 male,
Ontario, Ottawa 32 male, Maryland, Dorchester Co 33-35 A. floricola: 33 male, Ontario, Port Cun-
nington 34 male, Ontario (46.9300, -79.7268) 35 male, Ontario, Gravenhurst 36-38 A. inexpectus:
36, 37 male, Tuva (50.6690, 92.9844) 38 female, Tuva (51.316, 94.495). For additional images of the
floricola group, see Figs 49-58.

At stake is not the name used for the common white-striped species (which would
be floricola regardless), but the name for the uncommon dusty brown species, which
would be palustris were we to accept these specimens as its types. However, as argued
above, they are not the types. We therefore treat palustris as a synonym of floricola, and
sylvestris as the name for the dusty brown species. To settle the mislabelling properly, a
male specimen of the white-striped species from Wisconsin (the type locality) should
be designated as the neotype or lectotype of palustris. We have not yet done so as we

20 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

+
an 7
¥ ya yy

Attulus finschi
41

Attulus pubescens
43

Attulus terebratus
45

40

* Attulus fasciger

41,42 A. finschi: 41 male, Saskatchewan (55.31, -105.11) 42 male body, Ontario, 4 miles S of Wawa
44, 45 A. terebratus: 44 male, Novosibirsk Oblast (53.730, 77.865) 45 female, Novosibirsk 46, 47 A.
relictarius male, Stavropol Krai, (43.88, 42.70). For additional images of Attulus (Sitticus), see Figs 74-88.

await reexamination of the full Peckham collection in case specimens can be located
that might be identifiable as from the true type series.

Material examined. Canapa: BritisH CoLuMB1A: Richmond (2 females), 49.66,
-114.73 (1 female), 49.45, -115.08 (3 males, 6 females); ALBERTA: 52.46, -113.94
(1 male); Ontario: Richmond (2 males, 1 female), Gravenhurst (3 males), Port
Cunnington (1 female); Dwight (2 males, 5 females), Batchawana Bay (1 female),
Woodstock (3 females), 46.9300, -79.7268 (2 males, 1 female), 42.53, -80.12 (1 fe-
male), 43.2626, -80.5636 (1 male), 49.0852, -81.3237 (1 female); QuEBEc: Touraine
(1 male); Nova ScortA: 44.4318, -64.6075 (1 male); U.S.A.: WasHINGTON: 46.43,
-123.86 (2 males); CoLorapo: Jackson Lake State Rec. Area (1 male); NEBRASKa:
41.88, -103.09 (1 female).

Sitticine jumping spiders 2

Colonus hesperus 4472
Breda bicruciata 471
Aillutticina Aillutticus nitens 4475
Attinella dorsata d490
Jollas cupreus 4474, 4473
. ay Jollas leucoproctus 4478
Seay Jollas-Tomis
Sitticini 700 clade 90,100 Jollas cellulanus 4479
Sittisax ranieri 4481
100 Tomis palpalis 4477
Sitticina ra = Tomis manabita 4476
Pe Attulus (Sittilong) longipes ARV4504
Attulus (Sitticus) finschi 4480
Attulus Attulus (Sitticus) pubescens 4483
108 100 Attulus (Sitticus) terebratus RU18-5346
Attulus (Sitticus) fasciger 4487
Ta4 Say Attulus (Attulus) mirandus Ru 18-7308
Attulus (Attulus) saltator 4512
72.95 700 Attulus (Attulus) distinguendus RU18-6432
Attulus (Attulus) ammophilus d482
92,96 Aitulus (Attulus) zimmermanni RU18-5156
Aitulus (Attulus) burjaticus RU18-7302
100 Attulus (Attulus) inexpectus RU18-6799
100 —— Alttulus (Attulus) floricola 4438
Attulus (Attulus) rupicola 4491, d492

48 Attulus (Attulus) sylvestris 4489
1.

UCE phylogeny

Figure 48. Maximum likelihood phylogeny from 757 concatenated UCE loci (average 113231 base
pairs/taxon) analyzed primarily for the 23 Core Taxa in black (IQ-TREE, partitioned by locus). Topology
is identical in unpartitioned analyses, with nearly identical branch lengths. Bootstrap percentage values
from 1000 replicates shown for each clade. Where two numbers are shown, the first is the bootstrap
percentage for the partitioned analysis, the second for the unpartitioned analysis. Where one number
is shown, both analyses yielded the same percentage. An analysis of the All Taxa dataset, including the
weakly-sequenced taxa in grey, yielded the same topology.

Attulus (Attulus) sylvestris (Emerton, 1891), restored (removed from synonymy

with S. floricola)
Figures 31, 32, 54-58

Attus sylvestris Emerton, 1891 (Holotype male in MCZ from Beverly, Massachusetts,
examined).

Sitticus magnus Chamberlin & Ivie, 1944, syn. nov.

Sitticus rupicola — Prészynski, 1980, figs 58, 59 (misidentification), specimen from
Texas.

Remarks. A widespread but little-known Nearctic species, A. sylvestris can be found on
partially shaded ground where the males stand out for their tiny bouncing bright white
spots (the white tuft of setae on the palp’s tibia). We have found them on rocks and leaf
litter along a forest edge in Ontario, on the ground at the edge of a creek in a forest
in California, and on forest leaf litter in Maryland. See discussion under A. floricola

22 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

about why we judge A. sylvestris to be the proper name of this species, at issue because
of confusion over the type specimens of Attus palustris.

Both males and females have shorter legs and less contrasting markings than in A.
floricola, but the distinction of markings is most notable in the male, which lacks the
high-contrast white stripes on dark brown seen in A. floricola. The white setae on the
male’s palp are concentrated on just the tibia and end of the femur. The bulb of the palp
is rotated slightly more than in A. floricola, and thus the spermophore’s path shows an
upturn (i.e., the loop is angled to point distally instead of basally as in floricola), and the
female’s copulatory ducts arrive further to the posterior before looping back anteriorly
to enter the spermathecae. In these regards the genitalia resemble those of the Eura-
sian A. rupicola, A. caricis, and A. inexpectus (Logunov and Kronsestedt 1997). Aztulus
sylvestris is most similar to A. caricis in appearance (low-contrast brown markings), in
having a small loop of the copulatory duct, and small body size, but differs in brighter
markings on the palp, a more anteriorly-placed junction where the ducts enter the sper-
matheca, a larger epigynal RTA coupling pocket, and a more distinctly swollen bulb
of the spermatheca. (They are also distinct on the COI tree, Fig. 104.) The synonymy
of magnus can be determined by its original description and Prdszynski’s (1980) excel-
lent drawing of the vulva of the holotype female. The female from Texas tentatively
identified by Prdszynski (1980: 15, figs 58, 59) as S. rupicola is considered here to be S.
sylvestris based on his clear drawings showing the loop of the copulatory duct slightly
bigger than typical, but not reaching nearly as far to the posterior as in S. rupicola.

Material examined (all in UBC-SEM except as indicated): CANADA: ONTARIO: Ot-
tawa, Britannia Bay, 45.374, -75.796 (26 males, 3 females), Long Point, 42.53, -80.12
(2 females); U.S.A.: MaryLANb: Dorchester Co. (1 male 1 female, MCZ); CoLorapo:
Morgan Co., Fort Morgan (1 female); Cattrornia: Smith Redwoods State Reserve (1
male), 36.3907, -121.5951 (2 females), 36.3742, -121.5614 (1 male, 4 females).

Attulus (Attulus) striatus (Emerton, 1911)
Figures 59-63

Sitticus striatus Emerton, 1911

Remarks. Aitulus striatus is a small-bodied Northern species with distinctively striped
males, from sphagnum bogs. Although we were unable to obtain molecular data for it
or the similar A. rivalis and A. cutleri, these three species can be placed into subgenus
Attulus with some confidence, based on their boxy carapaces (resembling the other
Attulus (Attulus) rather than Attulus (Sitticus)), and the genitalic similarities with sub-
genus Aztulus, including the two small posterior openings of the epigyne on either side
of a narrow triangular RTA coupling pocket. Prészynski (1980) considered them close
to the floricola group in particular.

We reinstate S. rivalis Simon, 1937 as a distinct species (contra Prészynski 2017a),
accepting Logunov’s (2004) clear evidence for their distinction (primarily, in the rotation
of the bulb of the palp). Aztulus rivalis is known from France, also from sphagnum bogs.

Sitticine jumping spiders 23

Attulus floricola Attulus sylvestris Attulus striatus Attulus cutleri

aa,

é : 4 am * : ; .

—68. Sitticines of Canada: Attulus subgenus Attulus (for A. ammophilus, see Figs 69-73)
49-53 Aztulus floricola: 49 palp (Ontario, Gravenhurst) 50, 51 ventral view of epigyne, dorsal view
of cleared vulva (Ontario, Gravenhurst) 52 male (Ontario, 46.9300, -79.7268) 53 female (Ontario,
46.9300, -79.7268) 54-58 Aztulus sylvestris: 54 palp (Ontario, Ottawa) 55, 56 ventral view of epigyne,
dorsal view of cleared vulva (Ontario, Ottawa) 57 male (California, 36.3646, -121.5544) 58 female
(Ontario, 42.55, -80.13) 59-63 Azzulus striatus: 59 palp (Ontario, 45.1453, -75.8467) 60, 61 ventral
view of epigyne, dorsal view of cleared vulva (Ontario, 45.1453, -75.8467) 62 male (Ontario, 45.1453,
-75.8467) 63 female (New Hampshire, Ponemah Bog) 64-68 Aztulus cutleri: 64 palp (Northwest Ter-
ritories, Wrigley) 65, 66 ventral view of epigyne, dorsal view of cleared vulva (Northwest Territories,
Wrigley) 67 male (Northwest Territories, Inuvik) 68 female (Yukon, 67.57, -139.67). For habitus of
other Aztulus species, see Figs 15-38.

Pieter

‘< or

Figures 49

24 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

Material examined (all UBC-SEM): Canada: AtBerta: S. Islay (3 female), Bea-
verhill Lake (1 female); ONTARIO: 48.3260, -76.8365 (1 female); 3 km S. Richmond
(6 males, 2 females); NEw Brunswick: Chipman (1 male, 1 female); U.S.A.: New
Hampsuire: Ponemah Bog (1 female).

Attulus (Attulus) cutleri (Prészynski, 1980)
Figures 64-68

Sitticus cutleri Prdszytiski, 1980
Sitticus gertschi Prészynski, 1980

Remarks. A Sibero-American boreal species that is little collected, resembling closely
A, striatus but differing in having less striped legs, a less rotated bulb of the male palp,
more medially placed epigynal openings. Collected on “leaf litter under small Salix just
above stream” (D. Maddison, June 1981, Inuvik).

Material examined. Canapa: NorrHwest TERRITORIES: Wrigley (1 female,
CNC), Inuvik (1 male, UBC-SEM).

Subgenus Sitticus Simon, 1901
Figures 39-47, 74-88

Sitticus Simon, 1901 (type species Avaneus terebratus Clerck 1757)
Hypositticus Lohmander, 1944 (type species Aranea pubescens Fabricius, 1775)
Sittipub Prészynski, 2016 (type species Aranea pubescens Fabricius, 1775)

The species placed here, despite having palpi with very different embolus lengths, share
a similarly narrow and high body with relatively long legs (Figs 39-47), and (except
for A. relictarius) a dramatically large RTA, broadly arising from the tibia and sweeping
diagonally to the retrolateral and distal (Figs 74, 79, 84). Several species have a long
embolus and correspondingly long and convoluted copulatory ducts, though A. pube-
scens and A, relictarius have among the shortest in sitticines. The species of Sitticus are
Palearctic or Holarctic; the following three are found in Canada.

Attulus (Sitticus) finschi (L. Koch, 1879)
Figures 41, 42, 79-83

Attus finschii L. Koch, 1879
Euophrys cruciatus Emerton, 1891

Remarks. The natty contrasting black-and-white markings distinguish Aztulus finschi
from the closely related A. fasciger. Attulus finschi is the only Sitticus that has likely

Sitticine jumping spiders 29

been in the Americas for thousands of years; it also lives in Siberia. It is found in boreal
habitats on tree trunks.

Material examined (all UBC-SEM): Canapa: SaskaTCHEWAN: 55.31, -105.11
(1 male, 1 female), 55.27, -105.19 (1 female); ONTaRIo: Wawa (1 male), Nipigon (1
female), 48.9143, -80.9446 (2 females); New Brunswick: Doaktown (1 male).

Attulus (Sitticus) fasciger (Simon, 1880)
Figures 39, 40, 74-78

Attus fasciger Simon, 1880

Remarks. This species, introduced to North America apparently in the middle of the
20" century (Cutler 1990), is typically found on buildings. The large male palp and
spaghetti-like copulatory ducts distinguish it from other species in North America
except the differently-coloured A. finschi.

Material examined (all in UBC-SEM): Canapa: Ontario: Burlington (3 males,
6 females), 43.35074, -79.75928 (25 males, 14 females); U.S.A.: Missourr: Dogtown
(3 males, 4 females); MassacHuseTts: Cambridge (1 female).

Attulus (Sitticus) pubescens (Fabricius, 1775)
Figures 43, 84-88

Aranea pubescens Fabricius, 1775

Remarks. Although closely related to A. fasciger and A. terebratus, which have among
the longest emboli and copulatory ducts in sitticines, Attulus pubescens has among the
shortest known in sitticines. The very large RTA is distinctive. Introduced to North
America in the 20" century (Cutler 1990).

Material examined (All in UBC-SEM): Canapa: British CoLuMBiA: Vancouver
(1 male 1 female); U.S.A.: Massacnusetts: Cambridge (3 males, 3 females), Boston
(2 males), Milton (2 males), Arlington (1 female).

Subgenus Sittilong Proszynski, 2017
Sittilong Proszynski, 2017 (type species Attus longipes Canestrini, 1873)

The single species Attulus (Sittilong) longipes of the European Alps is peculiar for its
flat body and long first legs in the male, as well as its genitalia. Like Sittisax and other
members of the Jo/las- Tomis clade, the RTA is offset basally, and the epigynal openings
are anterior and medial. The little-studied Attulus niveosignatus has somewhat similar
genitalia and may also belong in Sittilong.

26 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

pS eras hg ETM ight Caaciee ye ; : :
Figures 69-88. Sitticines of Canada: Aitulus, continued 69-73 Azttulus (Attulus) ammophilus: 69 palp
(Ontario, Oakville) 70, 71 ventral view of epigyne, dorsal view of cleared vulva (Ontario, Hamilton)
72 male (British Columbia, 49.08, -119.52) 73 female (British Columbia, 49.08, -119.52) 74=78 A. (Sit-
ticus) fasciger (Ontario, 43.3508, -79.7593): 74 palp 75, 76 ventral view of epigyne, dorsal view of cleared
vulva 77 male 78 female 79-83 A. (S.) finschi: T9 palp (Ontario, Wawa) 80, 81 ventral view of epigyne,
dorsal view of cleared vulva (Saskatchewan, 55.31, -105.11) 82 male (Saskatchewan, 55.31, -105.11) 83 fe-
male (Saskatchewan, 55.27, -105.19) 84-88 A. (S.) pubescens: 84 palp (Massachusetts, Milton) 85, 86 ven-
tral view of epigyne, dorsal view of cleared vulva (Massachusetts, Arlington) 87 male (Massachusetts, Cam-

bridge) 88 female (Massachusetts, Cambridge). For other images of Aztulus (Sitticus), see Figs 39-47.

Sitticine jumping spiders 27
The Jollas-Tomis clade

We have chosen not to subdivide the Neotropical Sitticina more finely than into two
genera, Jomis and Jollas, primarily because the fauna is poorly enough known that it is
as yet unclear what coarseness of division would be most useful. We might have syn-
onymized their respective Nearctic offshoots (Sittisax into Tomis, and AZtinella into Jol-
las), but by retaining them as distinct, we facilitate the eventual splitting of both Zomis
and Jollas as their species become better known. We choose splitting in the Jollas- Tomis
clade, in contrast to lumping with Attulus, because the phylogenetic divergences are so
much deeper in the former compared to the latter.
The Jollas- Tomis clade includes four genera with 31 species:

Attinella Banks, 1905, with three species:
Attinella concolor (Banks, 1895), comb. nov., transferred from Sitticus
Attinella dorsata (Banks, 1895), combination restored, transferred from Sitticus
(type species)
Attinella juniperi (Gertsch & Riechert, 1976), comb. nov., transferred from Sittiab
Jollas Simon, 1901, with 12 species:
Jollas amazonicus Galiano, 1991
Jollas cellulanus (Galiano, 1989), comb. nov., transferred from Sitticus
Jollas cupreus W. Maddison, sp. nov.
Jollas flabellatus (Galiano, 1989), comb. nov., transferred from Sitticus
Jollas geniculatus Simon, 1901 (type species)
Jollas hawkeswoodi Makhan, 2007
Jollas leucoproctus (Mello-Leitao, 1944), comb. nov., transferred from Sitticus
Jollas manantiales Galiano, 1991
Jollas paranacito Galiano, 1991
Jollas pompatus (Peckham & Peckham, 1894)
Jollas puntalara Galiano, 1991
Jollas richardwellsi Makhan, 2009
Sittisax Prészynski, 2017, with two species:
Sittisax ranieri (Peckham & Peckham, 1909)
Sittisax saxicola (C. L. Koch, 1846) (type species)
Tomis F.O. Pickard-Cambridge, 1901, with 14 species
Tomis canus Galiano, 1977, combination restored, transferred from Sitticus
Tomis kratochvili (Caporiacco, 1947), comb. nov., transferred from Pseudattulus
Tomis manabita W. Maddison, sp. nov.
Tomis mazorcanus (Chamberlin, 1920), comb. nov., transferred from Sitticus
Tomis mona (Bryant, 1947), comb. nov., transferred from Sidusa
Tomis palpalis F. O. Pickard-Cambridge, 1901, combination restored, transferred
from Sitticus (type species)
Tomis pavidus (Bryant, 1942), comb. nov., transferred from Sidusa
Tomis phaleratus (Galiano & Baert, 1990), comb. nov., transferred from Sitticus
Tomis pintanus (Edwards & Baert, 2018, comb. nov., transferred from Sitticus

28 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

Tomis tenebricus (Galiano & Baert, 1990), comb. nov., transferred from Sitticus
Tomis trisetosus (Edwards & Baert, 2018), comb. nov., transferred from Sitticus
Tomis uber (Galiano & Baert, 1990), comb. nov., transferred from Sitticus
Tomis vanvolsemorum (Baert, 2011), comb. nov., transferred from Sitticus
Tomis welchi (Gertsch & Mulaik, 1936), comb. nov., transferred from Sitticus

Genus Attinella Banks, 1905, restored (removed from synonymy with Sitticus)

Attinella Banks, 1905 (type species Attus dorsatus Banks, 1895)
Sittiab Prészynski, 2017 (type species Sitticus absolutus Gertsch & Mulaik, 1936), syn. nov.

Small species from southern North America, related to the Jollas of South America.
Except for the thin longitudinal stripes of A. dorsata, their bodies are more or less un-
marked. Like many other members of the Jollas- Tomis clade, the RTA is long and thin,
paralleling the axis of the palp, the tibia is robust, and the embolus is fairly short. The
first leg’s trochanter is unusually long in at least some males (note angles in Fig. 12),
though less so than in Jo/las. The epigynal openings are anterior, with the ducts (intially
fused) leading to the posterior and to fairly small spermathecae. As noted below under
A, dorsata, the synonymy of Sitticus absolutus with Attus dorsatus leads to Sittiab being
a junior synonym of Aftinella. Two species of Attinella reach Canada.

Attinella concolor (Banks, 1895)
Figures 89—93

Attus concolor Banks, 1895 (holotype examined; see Maddison 1996: 270)
Sittacus cursor Barrows, 1919, synonymy restored

Sitticus floridanus Gertsch & Mulaik, 1936

Remarks. A small unmarked leaf litter species, known best from the southeastern Unit-
ed States, but recently reported from Canada in the BOLD barcode database (Ratnas-
ingam and Hebert 2007, 2013), from the extreme southern point in Ontario (Point
Pelee National Park, specimens PPELE142-11, PPELE183-11, CNPPE2332-12,
PPELE666-11, PPELE644-11).

Prészynski (2017a) rejected Maddison’s (1996) synonymy of cursor with concolor
on the basis of “lack of documentation”, an extra specimen inside the type vial, and
the fact that it was published in a revision of Pelegrina. However, Maddison (1996)
indicated clearly the evidence that identified the holotype within the vial (by its loca-
tion, labeling, and match to Banks's description), and the features that matched the
specimen to Sittacus cursor Barrows; that the nomenclatural act was published in a revi-
sion of a different salticid genus has no bearing on the validity of the act. Maddison’s
synonymy, therefore, is reaffirmed here as valid.

Material examined. U.S.A.: FLortpa: Gainesville (1 male, 1 female, UBC-SEM).

Sitticine jumping spiders 29

91 r 96 101
Attinella concolor Aitinella dorsata Sittisax ranieri
92 } 97
,

Figures 89-103. Sitticines of Canada: the Jollas- Tomis clade, represented by the genera Attinella and Sit-
tisax 89-93 Attinella concolor: 89 palp (Florida, Gainesville) 90, 91 ventral view of epigyne, dorsal view
of cleared vulva (Florida, Gainesville) 92 male (Texas, 30.10, -97.25) 93 female (Texas, 30.10, -97.25)
94-98 Attinella dorsata: 94 palp (California, San Diego County) 95, 96 ventral view of epigyne, dorsal
view of cleared vulva (British Columbia, Nanaimo) 97 male (California, Siskiyou County) 98 female
(British Columbia, 48.870, -123.379) 99-103 Sittisax ranieri: 99 palp (Northwest Territories, Tuktoyak-
tuk) 100, 101 ventral view of epigyne, dorsal view of cleared vulva (Nunavut, Baffin Island) 102 male
(Saskatchewan, 55.27, -105.19) 103 female (Ontario, Old Woman Bay).

30 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

Attinella dorsata (Banks, 1895)
Figures 11-14, 94-98, 105

Attus dorsatus Banks, 1895 (holotype female in MCZ from California: Los Angeles,
examined)

Sitticus absolutus Gertsch & Mulaik, 1936, synonymy restored

Sitticus callidus Gertsch & Mulaik, 1936, synonymy restored

Remarks. While females of this small Southwestern desert-dwelling species are indis-
tinctly unmarked, males tend to be reddish with a narrow central longitudinal stripe
(Figs 11-14). Prdészyfiski (2017a) rejected Richman’s (1979) synonymy of Attinella dor-
sata (Banks, 1895) with Sitticus absolutus, saying that dorsata is unidentifiable. That
statement is false, given that the type specimen is in the MCZ and in good condition.
The specimen (examined) has a relatively wide carapace with single thin longitudinal
pale line dorsally, long fourth leg, no retromarginal cheliceral tooth, and epigyne (Fig.
105) with a single anterior opening that leads posteriorly through a single duct that
splits before the spermathecae, which are visible as two small medial pear-shapes flanked
by slightly larger chambers. In these respects, it clearly falls within our current con-
cept of Sitticus absolutus as a common, widespread, and relatively uniform species from
Texas to California north to Canada (see illustrations by Gertsch and Mulaik 1936,
Prészynski 1973). Even if future work were to show that the Californian populations
(type locality of dorsatus) and Texan populations (type locality of absolutus) represent
distinct species, they are extremely closely related, certainly congeneric. Attus dorsatus is
a member of these Californian populations, and for this reason the synonymy of Sittiab
(type species Sitticus absolutus) with Attinella (type species Attus dorsatus) is assured.

Material examined. Canapa: BritisH CoLuMBIA: Summerland (1 male, CNC),
Galiano Island (2 males, 3 females, UBC-SEM), Nanaimo (1 female). U.S.A.: CALI-
FORNIA: Humboldt Co., Orleans (1 male, UBC-SEM), Siskyou Co., Beaver Creek
and Klamath River (1 male, UBC-SEM), San Diego Co., Johnson Canyon (1 male 1
female, UBC-SEM), El Dorado Co., Camino (1 female, UBC-SEM), Inyo Co., Gil-
bert Summit (1 female, UBC-SEM); Uran: Millard Co., Sevier Lake (1 male, UBC-
SEM); Cotorapo: Morgan Co., Jackson Lake (1 male, UBC-SEM), Jefferson Co.,
Golden (2 females, UBC-SEM); Texas: Jim Hogg Co., Guerra (1 female, UBC-SEM),
Pecos Co., Fort Stockton (1 female, UBC-SEM).

Genus Jollas Simon, 1901
Figures 7-10, 108-119

Jollas Simon, 1901 (type species Jollas geniculatus Simon, 1901)
Oningis Simon, 1901 (type species Neon pompatus Peckham & Peckham, 1893)

Sitticine jumping spiders 31

Attulus sylvestris (Ontario: 41.9313, -82.5133) JN308621

Attulus sylvestris (Ontario: 41.949, -82.521) KM832264

Attulus sylvestris (California: 36.3646,-121.5544) d489

Attulus caricis (Netherlands: Overijssel) RMNH.ARA.12587
Attulus rupicola (Switzerland: 46.516, 9.65) KX039377
Attulus floricola (Germany: 53.71, 8.80) KY269335

Attulus floricola (Germany: 51.8792, 12.349) KX537297
Attulus floricola (Germany: Roehricht) KX537154

Attulus floricola (Saskatchewan: 49.153, -107.517) HQ928167
Attulus floricola (Yukon: 60.748, -137.513) MF815131
Attulus floricola (Alaska: 64.867, -147.74) KU876202

Attulus floricola (British Columbia: 51.4429, -116.542) JF 884697
Attulus floricola (Ontario: 43.505, -80.2092) GU682461

Attulus floricola (Northwest Territories: 60.023, -112.95) KP654279
Attulus floricola (Manitoba: 58.76, -94.086) GU684425

Attulus floricola (Saskatchewan: 49.071, -106.526) MF816431
Attulus floricola (Alberta: 53.192, -117.956) KM825931

Attulus floricola (Saskatchewan: 52.4898, -107.3843) d488
Attulus floricola (New Brunswick: 45.625, -65.06) MF815448
Attulus floricola (Alberta: 53.674, -112.831) KP656704

Attulus floricola (New Brunswick: 46.8161, -64.915) MF814227

Placement of COl-barcoded specimens
of the floricola-group

104

Figure 104. Relationships among Attulus floricola mitochondrial COI sequences in the context of the
floricola group. Specimens in bold had their relationships constrained by the UCE phylogeny of Fig. 48;
not shown are the relationships outside the floricola group, which are fixed to match the UCE phylogeny.
The placement of non-bold specimens on this constrained skeletal tree was inferred by maximum likeli-

hood (RAxML, codon positions as separate partitions).

———$—$ $$ rn

Attinella dorsata Tomis welchi

Figures 105-107. Epigynes of Aztinella dorsata and Tomis welchi \05 holotype of Attus dorsatus Banks,
1895, epigyne, ventral view 106, 107 holotype of Sitticus welchi Gertsch & Mulaik, 1936 106 epigyne,

ventral view 107 cleared vulva, dorsal view.

A Neotropical group, consisting of two species groups, the small glabrous or shiny
geniculatus group (Galiano 1991b), and the typically grey /eucoproctus group (Galiano
1989). The male’s first trochanter is relatively long, approximately as long as the coxa
(Galiano 1989). Typically, the male’s first tibia and patella are marked by dark lines
on the prolateral face. Epigynal openings are medial, with ducts proceeding toward
the lateral. Most species have a long thin RTA, though that is also seen in many Tomis
and Attinella.

32 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

Jollas cupreus W. Maddison, sp. nov.
http://zoobank.org/68F87DD6-8C3 1-4D0B-A349-245B9B201CF3
Figures 7, 108-111, 113-119

Type material. Male holotype and 2 male, 3 female paratypes from Ecuapor: Orel-
lana: Rio Bigal Reserve, main camp area. 0.5251, -77.4177. 950 m elev. 1-5 No-
vember 2010. W & D Maddison, M Vega, M Reyes. WPM#10-041c. The holotype
(specimen ECU2010-2060) pertains to the Museum of Zoology, Pontificia Uni-
versidad Catdlica, Quito, Ecuador (QCAZ), but is currently held in the Spencer
Entomological Collection at the Beaty Biodiversity Museum, University of British
Columbia (UBC-SEM).

Etymology. Refers to the copper colour of males.

A species common in eastern Ecuador on disturbed open grassy ground. It was
used in the molecular phylogenetic study of Maddison and Hedin (2003) under the
name “Jollas sp.” (voucher $162) from Sucumbios, Ecuador.

Diagnosis. Differs from the very similar Jollas puntalara Galiano, 1991 in the thin-
ner and straighter RTA and the angle at which the embolus arises. The RTA is more
or less straight until a curl at the tip, but it narrows dramatically for its terminal three
quarters (Fig. 109), whereas in J. puntalara (Galiano, 1991b: fig. 26) it bends at the
midpoint and thins much less dramatically. The embolus of /. cupreus, as it arises, pro-
ceeds directly to the prolateral, thus generating an angle in the retrolateral-basal corner
of the bulb (like a chin pointing to the retrolateral), while the embolus of J. puntalara
emerges angled toward the basal, leaving the bulb more rounded (arrow in Fig. 112).
These differences are small but consistent, insofar as all Ecuadorian specimens show
the distinct “chin” at the base of the embolus and the narrower RTA. It might usually
be conservative to leave such close forms as a single species, but given that there is con-
siderable data (molecular phylogenetic and chromosome) attached to the Ecuadorian
form, it is safer to name it and thus provide an unambiguous anchor for these data.
(Cristian Grismado kindly supplied photographs of Galiano’s (1991b) holotype of Jol
las puntalara to facilitate our comparison, although these differences can be seen as well
in her figures 26 and 29.)

Description. Male (holotype). Carapace length 1.37; abdomen length 1.16. Car-
apace orange with a black ocular area, mostly glabrous, with only a few scattered se-
tae. Clypeus orange-brown. Chelicerae vertical, orange. Palp orange-brown except for
dark brown cymbium, with dark setae except brilliant white patch of setae dorsally on
patella. Legs long, especially the first and fourth. Legs honey coloured to orange-brown
except for a strong black line on prolateral-ventral face of first patella, tibia and meta-
tarsus. Embolus arises at ca. 5 o'clock and curls half-way around bulb. Tibia somewhat
bulbous, broad, with bases of setae on retrolateral side forming row of tubercles. RTA
begins broad but then narrows abruptly at ca. one quarter its length, from which point
it proceeds straight until just before the tip, where it curls. Abdomen orange-brown,
with black scalloped patch covering dorsum, covered with metallic scales. A patch of
bright white setae sits above the anal tubercle.

Sitticine jumping spiders bb)

108 ventral view 109 retrolateral view 110 ventral view of epigyne of paratype I 11 dorsal view of same,

cleared 112 palp of holotype of /. puntalara Galiano 113-115 holotype male 116 male from Yasuni,
Ecuador (-0.675, -76.397) | 17 holotype male in alcohol 118, 119 paratype female.

Female (paratype). Carapace length 1.36; abdomen length 1.89. Much darker
than the male in body and appendages (Figs 130, 131). Carapace dark brown, black
in ocular area, sparsely covered with paler scales. Clypeus and chelicerae brown, more
or less glabrous. Chelicerae brown, more or less. Palps and legs honey coloured but
with strongly contrasting black markings: annulae at joints, black stripes or patches on
front and back faces of femora, and black stripe on front face of first and second tibiae.
Abdomen black but with reflective metallic scales. Epigyne (Figs 110, 111) with dis-
tinctive dark inverted “V” in which are the narrow openings into the copulatory ducts,
though lateral pockets may lead the embolus to the openings.

Additional material. 22 males and 6 females from: Ecuapor: Napvo: Tarapoa. 23
June — 1 July 1988 W. Maddison WPM#88-002 (1 male); Ecuapor: Napo: bridge

34 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

over Rio Cuyabeno on road to Tipishca. 25-30 June 1988 W. Maddison WPM#88-
004 (1 male 1 female); Ecuapor: Napo: bridge over Rio Cuyabeno on road to Tip-
ishca. 29-30 July 1988 W. Maddison WPM#88-018 (4 males 2 females); Ecua-
por: Napo: Reserva Faunistica de Cuyabeno, Laguna Grande, Sendero La Hormiga.
2—5 August 1988 W. Maddison WPM#88-023 (2 males); Ecuapor: Napo: Reserva
Faunistica de Cuyabeno, Laguna Grande, PUCE field station. 1-7 August 1988 W.
Maddison WPM#88-025 (1 male); Ecuapor: Napo: bridge over Rio Cuyabeno on
road from Lago Agrio to Tipishca. 8—9 August 1988 W. Maddison WPM#88-027
(1 male); Ecuapor: Sucumsios: Reserva Faunistica Cuyabeno, Laguna Grande,
PUCE field station. 0.002, -76.172. 21-29 July 1989 W. Maddison WPM#89-032
(1 male); Ecuapor: Sucumsios: bridge over Rio Cuyabeno on road between Tara-
poa and Tipishca, 0.025, -77.308. 29 July 1989 W. Maddison WPM#89-036 (1
male); Ecuapor: SucumBios: Reserva Faunistica Cuyabeno, Nuevo Mundo cabins
along Rio Cuyabeno at jcn with Lago Agrio-Tipishca HWY 19-29 April 1994 W.
Maddison WPM#94-021 (3 males); Ecuapor: SucumBios: Reserva Faunistica Cuy-
abeno, Nuevo Mundo cabins, jcn Rio Cuyabeno & Lago Agrio-Tipishca HWY tree
trunks 19-29 April 1994 W. Maddison WPM#94-023 (1 male); Ecuapor: Mo-
RONA SANTIAGO: km 3 from Limén towards Gualaceo. 2.9663, -78.4209; 1250 m
el. 12 July 2004 Maddison, Agnarsson, Iturralde, Salazar. WPM#04-030 (1 male
2 females); Ecuapor: Morona SantTiaco: km 4 from Limén towards Gualaceo.
2.9808, -78.4414; 1380 m el. 12 July 2004 Maddison, Agnarsson, Iturralde, Sa-
lazar. WPM#04-031 (2 males); Ecuapor: OrRELLANA: Yasuni Res.Stn.area, Station
area 0.675, -76.397 210-280 m elev. 26 July — 13 Aug 2011 Maddison/Piascik/
Vega WPM#11-015 (2 males); Ecuapor: OrELLANA: Yasuni Res.Stn.area, Station
area 0.674, -76.397 210-280 m elev. Clearings, forest edge 8—9.Aug.2011 Maddi-
son/ Piascik/Vega. WPM#11-104 (1 male); Ecuapor: OrRELLANa: Rio Bigal Reserve,
boundary along road. 0.541, -77.424. 970 m elev. 5 November 2010. M Vega, D &
W Maddison, M Reyes. WPM#10-048 (1 female). (Note: the province Sucumbios
was established after 1988; the 1988 localities listed as Napo Province would now all
be in Sucumbios.).

Genus Sittisax Prészynski, 2017, restored (removed from synonymy with Sitticus)

Sittisax Prészynski, 2017 (type species Euophrys saxicola C.L. Koch, 1846)

Breitling’s (2019) synonymy of Sittisax with Sitticus is here rejected based on our phy-
logenetic results, which strongly support it as the sister group of Zomis. According to
the phylogeny, this lineage of two species arrived from the New World to Eurasia inde-
pendently from Aztulus, and retains a few features more similar to the other members
of the Jollas- Tomis clade: the RTA is offset basally, and the epigynal openings are placed

anteriorly and medially.

Sitticine jumping spiders 35

Sittisax ranieri (Peckham & Peckham, 1909)
Figures 99-103

Attus lineolatus Grube, 1861 (junior homonym)
Sittacus ranieri Peckham & Peckham, 1909

Remarks. ‘The Holarctic Sittisax ranieri is a widespread boreal species, which in North
America follows the high elevations of the Western Cordillera to the south, living on
rocks and litter. It is dark in colour, large-bodied, and with distinctive genitalia.

Material examined. Canapa: NorTHWEST TERRITORIES: Tuktoyaktuk (1 male);
Nunavut: Baffin Island (1 female); Britis CoLtumsia: Downton Creek (2 males 2
females), 49.026, -114.061 (1 male), 59.8, -127.5 (1 male), Pink Mountain (1 male);
YuKON: km 72 Dempster Highway (2 males, 2 females); km 75.6 Dempster Highway
(1 female); SASKATCHEWAN: 55.27, -105.19 (2 males), ONTARIO: Old Woman Bay (1
female); New BRUNSWICK: 65.336, -69.4 (6 males, 3 females); U.S.A.: WASHINGTON:
Spray Park (1 males, 2 females); OREGON: 45.261, -117.178 (1 female); CoLorapo:
39.803, -105.782 (1 male).

Genus Tomis K.O. Pickard-Cambridge, 1901, restored (removed from synonymy
with Sitticus)
Figures 5, 6, 106, 107, 120-128

Tomis F.O. Pickard-Cambridge, 1901 (type species Tomis palpalis FO. Pickard-Cam-
bridge, 1901)

Pseudattulus Caporiacco, 1947 (type species Pseudattulus kratochvili Caporiacco,
1947), syn. nov.

A Neotropical group whose male spermophore (with some possible exceptions) has
a “shortcut loop”. That is, the large loop of the spermophore that normally occupies
much of the visible face of the tegulum, and which points basally in many sitticines
(e.g., Fig. 89), is incomplete, instead diving into the subtegulum, and thus not return-
ing terminally to complete the loop on the surface of the tegulum (e.g., Fig. 120;
Galiano 199 1a: fig. 13).

The phylogeny strongly places 7’ palpalis, T: manabita, and Sittisax ranieri togeth-
er. Although the phylogeny gives us the freedom to synonymize Sittisax into Tomis,
this deep clade will eventually deserve at least two genera, and so we tentatively retain
the boundary between the Neotropical 7omis and the Holarctic Sittisax, based on the
apparent difference in spermophore loops. The other species are included in Tomis
because of their apparent relationship with 7 palpalis and T’ manabita. The palpalis
group (T" palpalis, T: canus, 1: mazorcanus, 1; phaleratus, T: vanvolsemorum, and T:
uber) is delimited by a flattened cymbium (Galiano, 1991a) and well-separated epigy-

36 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

nal openings. The remaining species all are known from coastal areas of the Caribbean
or South America, and at least some live on beaches. ‘They might not form a monophy-
letic group, as some show a long thin RTA, others not. 7’ pavidus and T’ mona appear
especially close to 77 manabita by similarities in the palps. The others can be tentatively
included in Yomis because they share with 7’ palpalis and T: manabita the shortcut
spermophore loop.

The placement of Sitticus welchi Gertsch & Mulaik, 1936 in Tomis is tentative. The
holotype female (AMNH, examined) lacks most of its legs and setae, but is nonethe-
less identifiable as a sitticine through the absence of a retromarginal cheliceral teeth
and a very long leg that appears to be (it is disarticulated) of the fourth pair. The single
anteriorly-placed epigynal opening (Figs 106, 107) indicates it belongs in the Jollas-
Tomis clade. What suggests placement in Yomis in particular is the deep incision from
the epigastric furrow toward the epigynal opening (Fig. 106). Such an incision as seen
also in TYomis mona (Bryant 1947: fig. 6), which itself is placed in Tomis by the close
similarity between its palp and that of 77 manabita.

We synonymize Pseudattulus (see Ruiz et al. 2007) based on its shortcut spermo-
phore loop and flattened cymbium, which suggest close relationship to (or member-
ship in) the palpalis group. We accept (and thus re-assert) Ruiz et al.’s (2007) syn-
onymy of Sitticus cabellensis Proszyfski, 1971 with Pseudattulus kratochvili. Proszyhski
(2017a) had rejected their synonymy, but we see no basis for this, as Ruiz et al. had
explained it well. Although we suspect Pseudattulus will eventually be reinstated, keep-
ing it separate now would most likely yield a non-monophyletic genus Yomis. For
many species (e.g., those from the Galapagos) we have no basis for choosing whether
to assign them to Pseudattulus or to Tomis, and so either or both genera, if separated,
would likely be non-monophyletic. Uniting them solves this until we have better phy-
logenetic information.

Tomis manabita W. Maddison, sp. nov.
http://zoobank.org/4C656386-8B 15-4B5C-BF3F-27805897C65F
Figures 120-128

Type material. Male holotype, 10 male and 8 female paratypes from Ecuapor:
Manasi: Puerto Rico, Cabafias Alandaluz 5 May 1994 W. Maddison WPM#94-031.
The holotype (specimen UBC-SEM AR00217) pertains to the Museum of Zoology,
Pontificia Universidad Catélica, Quito, Ecuador (QCAZ), but is currently held in the
Spencer Entomological Collection at the Beaty Biodiversity Museum, University of
British Columbia (UBC-SEM).

Etymology. Based on the type locality; the form is the adjective in Spanish (mas-
culine or feminine).

A species on the beaches of coastal Ecuador, resembling Aztu/us in habitus. It was
used in the molecular phylogenetic study of Maddison and Hedin (2003) under the
name “Sitticus sp.” (voucher S220) from Manabi, Ecuador.

Sitticine jumping spiders af

Figures |20-128. Zomis manabita, sp. nov. 120, 121 Left palp of holotype 120 ventral view 121 retro-
lateral view 122 ventral view of epigyne of paratype 123 dorsal view of same, cleared 124—128 specimens

from type locality 124 male 125 male 126 female 127 male holotype 128 female paratype.

Diagnosis. Palp closely resembles that of 7omis pavidus, from which it differs in
the smaller tibia and longer RTA.

Description. Male (holotype). Carapace length 1.58; abdomen length 1.51. Car-
apace (Figs 124, 127) medium brown with recumbent brown setae except for thin me-
dial longitudinal band of white setae on thorax, two spots of pale setae in ocular area,
and a marginal band of white setae that is broad on the thorax but narrows to the an-
terior, disappearing before the clypeus is reached. Clypeus brown, with a few brownish
setae. Chelicerae vertical, with a few pale setae near the clypeus. Retromarginal cheli-
ceral teeth lacking. Palp clothed with white setae dorsally, but prolaterally with darker
integument and setae from tip of femur to cymbium; cymbium mostly dark brown.
Embolus (Fig. 120) arises broadly, more centrally beneath bulb (and less peripherally)
than is typical, then narrows abruptly at ca. 9 o'clock. RTA extremely long and thin,
parallel to axis of the palp. Legs honey-coloured, with notably darker annulae at the

38 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

tarsus-metatarsus joints, and black stripe on prolateral-ventral face of first patella and
tibia. Abdomen (Figs 124, 127) brown with two lateral and one median longitudinal
bands of paler setae, the medial band less distinct, wavy, and accompanied by a lateral
extension that forms a cross.

Female (paratype # UBC-SEM AR00218). Carapace length 1.76; abdomen
length 2.22. Carapace (Figs 126, 128) brown, covered unevenly with recumbent
cream-coloured setae. Clypeus with white setae, longest at midline where they over-
hang the chelicerae. Chelicerae brown, with a few setae near clypeus. Palps and legs
honey coloured, with weak annulae. Abdomen brown, marked as in male except bands
are less distinct (Figs 126, 128). Epigynum with an anterior atrium from which short
copulatory ducts lead diagonally to the spermathecae (Figs 122, 123).

Additional material. 15 males and 7 females from Ecuapor: Manast: Machalilla
National Park, Salaite, between HWY and coast 6 May 1994 W. Maddison WPM#94-
032 (4 males, 2 females); Ecuapor: Manast: Machalilla National Park, Salaite, 1 km
inland along trail from HWY. 6 May 1994 W. Maddison WPM#94-033 (3 males);
Ecuapor: Manast: Machalilla National Park, trail between Agua Blanca & San Se-
bastien 50-400 m dry forest 7 May 1994 W. Maddison WPM#94-034 (1 male); Ec-
uADOR: Manasi: Crucita. 30 August 1988 W. Maddison WPM#88-040 (2 males 4
females); Ecuapor: Det Oro: Jambeli 13 August 1989 W. Maddison WPM#89-040
(3 males); Ecuapor: Manasf: Puerto Lopez. 1.5497, -80.8104; 5 m el. 1-5 August
2004 W. Maddison. WPM#04-067 (2 males 1 female).

Species misplaced as sitticines

The following species are not sitticines, indicated by the presence of retromarginal chelicer-
al teeth (lacking in the Sitticini, a synapomorphy) or characteristic euophryine genitalia.

The following three are members of the Euophryini. They are left within sitticine
genera because it is unclear to which genus they should be transferred.

Jollas armatus (Bryant, 1943)

Jollas crassus (Bryant, 1943)

Jollas minutus (Petrunkevitch, 1930)

The following two species described in Sitticus are also euophryines (see Prdészynski
2017a). They are tentatively placed in a likely genus, Chinophrys:

Chinophrys taiwanensis (Peng & Li, 2002), comb. nov.

Chinophrys wuae Peng, (Tso & Li, 2002), comb. nov.

The following species can be moved out of Sitticus to their appropriate genera. The
type specimens of both, in the MCZ, have been examined.

Heliophanus designatus (Peckham & Peckham, 1903), comb. nov. — bears the strid-
ulatory apparatus characteristic of chrysillines (Maddison 1987), as well as the body
form, markings and epigynum typical of Heliophanus.

Mexigonus peninsulanus (Banks, 1898), comb. nov. — appears as a typical Mexigonus
with euophryine genitalia.

Sitticine jumping spiders by,

Chromosome diversity and evolution

Chromosome observations

Table 2 summarizes the chromosome complements of the 18 sitticines studied along
with those reported in the literature (Hackman 1948, Kumbicak et al. 2014). Except
in Attinella concolor, all autosomes are acrocentric. Eight species have the usual chro-
mosome complement for male salticids, 13 pairs of acrocentric autosomes and X,X,0
sex chromosomes. ‘Three taxa (A. burjaticus, A. floricola, A. finschi) have the standard
XX0 sex chromosomes but an extra pair of autosomes to make 28a+XaXa0. Of the
remaining species, six have neo-Y systems of varied forms, while the seventh, Attinel-
la concolor, has apparently completed a series of Roberstonian fusions to generate all
metacentrics and halve the chromosome number to male 14m + Xm0. The following
account of our observations, species by species, gives the basis for our interpretation of
chromosome complements.

Chromosomes of the Jollas- Tomis clade

Attinella concolor. 14m+Xm0 (Figs 129, 130): Nuclei of first meiotic metaphase show
clearly seven pairs of metacentric autosomes and one metacentric X chromosome
(Figs 129, 130). The metacentric autosomes appear strikingly different from the
usual acrocentrics typical of spiders. Most of the bivalents are held together by just
one arm at first metaphase, the other free.

Attinella dorsata: 26a+XaXa0 (uncertain). Scored as 26a+XaXa0 in notes from the
1980s, the slides are too faded and degraded for precise re-count, but re-examina-
tion shows they have at least 13 acrocentric bivalents, and what looks like XX0.
Although we might have abandoned the score entirely, we include it here to show
that it is at least similar to the typical salticid complement, and not at all what is
seen in the close relative Attinella concolor.

Jollas cupreus: 26a+XaXa0. One first division nucleus appears clearly as 13 autosomal
bivalents plus two acrocentric Xs, while two more show the typical Xs side by side.

Sittisax ranieri: 24a+XmXaYm (Figs 132-136). The distinctive chromosome comple-
ment is confirmed by many clear nuclei. The sex chromosomes (Figs 132-136)
appear as a rabbit’s head (the Y) with two ears (the Xs), one of which is floppy (a
metacentric with an unpaired arm). The two arms of the metacentric Y and one
of each X meet together at a single point, a junction of four arms. That the “head”
and “ears” segregate to opposite poles is confirmed by second metaphase counts
(nine nuclei with 12 acro.+1 meta.; five nuclei with 13 acro.+ 1 meta.). That the
“head” is a Y and the “ears” are Xs is indicated by counts of 26 acrocentrics and
two metacentrics in mitotic metaphase of two young females from Wawa, Canada
(47.79N, 84.90W) (2 complete, countable nuclei found in each female, scored in
1986; now faded). Unlike Habronattus (Maddison, 1982) and most other species

40

Table 2. Chromosome complements observed for males of 17-18 species of sitticines. The autosomal
counts represent diploid complement, and thus 26a means 13 pairs of acrocentric autosomes. In the
chromosome counts, a = acrocentric (one-armed), m = metacentric (two-armed). Exx. is the number of
specimens; nuc. is the number of nuclei showing the full chromosome complement; +nuc sex is the num-
ber of additional nuclei showing the sex chromosomes (though not clearly the autosomes). Uncertainties
about scoring, in particular about Attinella dorsata, Attulus burjaticus and the specimen labelled “Aztulus

Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

rupicola/floricola” are explained under Chromosome observations.

Species
Jollas-Tomis clade

Autosomes < Sex chrom. Y present

Locality

exx nuc +nuc sex

Attinella concolor 14m Xm0 no U.S.A.: Gainesville, 29.63, -82.37 1 6 2
A. dorsata 26a? XaXa0? 2 U.S.A.: Dillon Cr., 41.57, -123.54 3 11
Jollas cupreus 26a XaXa0 no Ecuador: Tarapoa, -0.12, -76.34 1 2 1
Sittisax ranieri 24a XmXaYm yes Canada: Leguil Creek, 59.8, -127.5 2 10 1
Canada: Inuvik, 68.31, -133.49 1 1
Canada: Wawa, 47.79, -84.90 2 8
U.S.A.: Mt. Monadnock, 42.87, -72.11 1 3
Sittisax saxicola 24a XaXaXaYm or yes Switzerland: Flims, 46.9, 9.2 3 14 10
XmYaYaYa
Tomis manabita 26a XaXa0 no Ecuador: Crucita, -0.9, -80.5 1 3 2
Attulus
Attulus (Attulus) 26a XaXa0 no Canada: Toronto, 43.65, -79.32 3 9 1
ammophilus Russia: Uvs Nuur, 50.6690, 92.9844 2 ii 6
A, (A.) burjaticus 2? (28a?) XaXa0 no Russia: Uvs Nuur, 50.677, 92.99 1 1 7
A. (A.) caricis 26a XaXa0 no 38.6, 34.8 (Kumbicak et al. 2014) =
A. (A.) cutleri 26a XaXaYa yes Canada: Inuvik, 68.35, -133.70 1 3 4
A, (A.) floricola 28a XaXa0 no Canada: Barrie, 44.43, -79.65 1 vf 7
U.S.A.: Naselle, 46.43, -123.86 1 2
A. (A.) rupicola/floricola 24a? XaXaXaYm? yes Switzerland: Flims, 46.9, 9.2 1 3 5
A, (A.) inexpectus 26a XaXa0 no Russia: Uvs Nuur, 50.6690, 92.9844 2 13 5
A. (A.) striatus 24a XaXaXaYm yes U.S.A.: Ponemah, 42.82, -71.58 1 5 6
Attulus (Sitticus) fasciger 26a XaXa0 Canada: Burlington, 43.351, -79.759 3 16
A. (S.) finschi 28a XaXa0 no Canada: Nipigon, 49.01, -88.16 1 4
Canada: Sault Ste. Marie, 46.94, -84.55 1 1
Canada: Chinook L., 49.67, -114.60 1 8 3
A, (S.) pubescens 26a XaXmYa yes U.S.A.: Cambridge, 42.38, -71.12 4 10 9
A. (S.) terebratus 26a XaXa0 no Russia: Karasuk, 53.730, 77.866 1 9 14
26a XaXa0 Hackman (1948) =

of sitticines, no heteropycnosis or achiasmate meiotic pairing was noted in the sex
chromosomes of S. ranieri which would have indicated ancestral X material. We
thus have no account for how this structure evolved, and what parts of it represent
ancestral X versus autosome material.

Sittisax saxicola: 24a+XaXaXaYm or 24a+XmYaYaYa (good quality, though ambiguous

in interpretation; Figs 137-139). The sex chromosome system in Sittisax saxicola
is at least superficially similar to that in S. ranieri except that the “rabbit” has three
straight “ears”. The many metaphase I nuclei observed show 12 clear autosomal
acrocentric bivalents plus the sex chromosomes, while two mitotic nuclei had clear
counts of 28 chromosomes, one of which is notably longer than the others, pos-
sibly the metacentric. The acrocentric “ears” of the sex chromosomes are always
oriented together toward one pole at metaphase I, the metacentric “head” to the
other, indicating either a XXXY or XYYY sex chromosome system. Consistent

Sitticine jumping spiders 4]

be i a i] z= —_

Figures 129-139. Chromosomes of first meiotic division in males of the Jollas- Tomis clade 129, 130 Az-
tinella concolor, with only seven pairs of autosomes, but each two-armed, 14m+Xm0, Florida (29.63N,
82.37W) 131 Tomis manabita, showing the two Xs off to one pole, and 13 acrocentric bivalents on the
metaphase plate, Ecuador (0.98, 80.5W) 132-136 Sittisax ranieri, whose distinctive XmXaYm appears
as a rabbit head with a droopy ear. White triangles show points where two bivalents are apparently linked
together 134-136 details of XXY of S. ranieri 137-139 Sittisax saxicola, with sex chromsomes, inter-
preted tentatively as XaXaXaYm, appearing as a rabbit head with three ears, Switzlerland (46.9N, 9.2E).

with this are three observations of second division nuclei with 15 acrocentrics, and
one observation with 12 acrocentrics and a metacentric. There is no clear evidence
from heteropycnosis, and no female karyotype, to indicate whether the “ears” are
the Xs or the Ys. We might invoke parsimony to suggest the metacentric is the Y
and the ears the Xs, as in S. vanieri, but will resist this, and treat the sex chromo-
somes as ambiguous, either XXXY or XYYY.

All four sex chromosomes of S. saxicola come together in a quintuple junction.
This and the quadruple junction of S. ranieri are unusual, possibly formed because

42 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

mutual translocations or repeated sequences generate a knit pattern of pairing.
White (1965) postulated that a similar triple terminal junction in a mantid is
formed by chiasmata joining the three arms on triple pairing segments and subse-
quently terminalizing. There is evidence that different autosomes in Sittisax might
also have common terminal segments. In all males of S. ranieri, autosomal biva-
lents with proximal chiasmata are often joined together into tetravalent and some-
times hexavalent chains, via the terminal ends of one chromosome of each bivalent
(see white triangles in Figs 132, 133). The terminal ends of the autosomes appear
to have small satellites.

Tomis manabita: 26a+XaXa0 (Fig. 131). Although there are only a few nuclei, they
show 13 autosomal bivalents plus two acrocentric Xs. In three nuclei, the two
acrocentric Xs are side by side and off to one pole.

Chromosomes of Attulus

Attulus (Attulus) ammophilus: 26a+XaXa0 (Figs 140, 141). Many clear nuclei show the
classical 13 acrocentric bivalents and two acrocentric X’s off toward one pole.
Attulus (Attulus) burjaticus: ?+XaXa0 (autosome count uncertain; Fig. 142). One clear
and isolated meiotic nucleus in metaphase I shows 15 figures, one of which is
presumably be the XX, suggesting that it may have 28a+XX0. Six nuclei show a
typical pair of XaXa toward one pole. The interpretation of XXO seems reasonably

secure, but the autosome count is not.

Attulus (Attulus) cutleri: 26a+XaXaYa (Figs 152, 153). There are a few clear nuclei, and
several more in which the sex chromosomes are clear (but the autosome counts
are not). Interpretation of the sex chromosomes seems fairly clear. They are inter-
preted to be XXY because two elements are seen side by side and slightly decon-
densed (the Xs). The third chromosome is small, paired terminally with the more
condensed end of the larger X, and thus interpreted as a Y. There is no hint of a
centromere in the larger X, and so all appear to be acrocentrics.

Attulus (Attulus) floricola: 28a+XaXa0, with one autosome much smaller (Figs 143-—
146). In addition to the clear division I nuclei showing the classic pair of X’s lying
side by side, counts of second division nuclei show either 14 acrocentric chromo-
somes (six clear nuclei) or 16 chromosomes (five clear nuclei). All of the second
division nuclei show one chromosome much smaller than the others. Those with
16 chromosomes show two of the chromosomes appearing larger and distinct in
appearance, consistent with their being the Xs, pointing to an XaXa0 sex chromo-
some system.

Attulus (Attulus) rupicola/floricola (Switzerland): 24a+XaXaXaYm (uncertain in details,
though the presence of at least one Y is secure; Figs 150, 151). The presence of a Y
chromosome is well supported, but the details of the sex chromosome system are
uncertain. No single nucleus shows both the chromosome count and the sex chro-
mosome system convincingly. The total number of chromosomes (27 acrocentrics
and one metacentric) can be seen in two mitotic nuclei, and in a few first division

Sitticine jumping spiders 43

Se UES RTS BW
= . *, | : 1 ; ae | .— “ .. 3
agi? (-7%..-3
te MS Age Bae Ee SD
ms “ ‘2 ov od ~ i
»g > sa + » . XaXa |
XaXa “A “>? s
es Y) Y ve @! ~ tid 4 - ;
w= : . a i » us os
fe Pages

Attulus ammophilus

ee e ; y 1

Figures 140-142. Chromosomes of first meiotic division of Attulus subgenus Attulus 140, 141 Aztulus
ammophilus, Tuva (50.6690N, 92.9844E): 140 four nuclei, three showing the two X chromosomes to-
ward one pole 141 two nuclei showing two Xs and thirteen pairs of acrocentric autosomes 142 Attulus
burjaticus, showing the two X chromosomes toward one pole, Tuva (50.677N, 92.99E). The three large

spots to the lower right are spermatids.

meioses. Although at least 20 nuclei show the V-shaped trivalent of metacentric
(point of the “V”) and two acrocentrics (distal arms of the “V”), interpreted as the
Y and two Xs, only three show the fourth member, an acrocentric, lying near one
of the Xs. This achiasmate association leads us to intepret the system as XaXaXaYm
rather than XmYaYaYa, but the evidence is weak, as there are no female counts,
heteropycnosis is not obvious, and most often the fourth member is lying distant
from the trivalent, usually not obviously directed to the same pole as the two acro-
centrics, though not apparently oriented against it either.

Attulus (Attulus) inexpectus: 26a+XaXa0 (Figs 147-149). Several very clear first division
nuclei show 13 acrocentric bivalents and the two acrocentric Xs, heteropycnotic
and lying side by side, off of the metaphase plate. Three second division counts are
consistent with an XX0 sex chromosome system (two counts of 13 acrocentrics;
one count of 15).

Attulus (Attulus) striatus: 24a+XaXaXaYm. The slides are too faded to score now even
under phase contrast, and so for this we rely entirely on notes from 1985. Those

44 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

Figures 143-153. Chromosomes of meiosis of Aztulus subgenus Attulus, continued 143-146 Aztulus
floricola, with an extra small bivalent (s) to make 28a+XaXa0, Ontario (44.43, -79.65): 143, 144 first
metaphase 145 second division, showing one nucleus with 14 acrocentrics, the other with 14 acrocentrics
and the two condensed Xs 147=149 Attulus inexpectus, showing 13 acrocentric bivalents and the sex chro-
mosomes (26a+XaXa0), Tuva (50.6690, 92.9844) 150, 151 Aztulus sp. (ambiguously identified, either A.
rupicola or floricola), tentatively intepreted as having 24a+XaXaXaYm, Switzerland (46.9, 9.2): 151 same,
sex chromosomes from another nucleus 152 Aztulus cutleri, with 26a+XaXaYa, Canada (68.35, -133.70)

153 same, sex chromosomes from another nucleus

Sitticine jumping spiders

Figures 154-164. Chromosomes of meiosis of Aztulus subgenus Sitticus 154 Attulus fasciger, three
nuclei, one showing the two Xs together and toward a pole, Canada (43.351N, 79.759W) 155-163 Az
tulus pubescens, with XaXmYa sex chromosomes, Massachusetts (42.38N, 71.12W) 157-161 XmYa sex
chromosomes from other nuclei; the second X is often not paired with them 162, 163 Second division

nuclei, all having 14 acrocentrics, and some having in addition a metacentric (m) 164 Aftulus terebratus,

two nuclei (26a+XaXa0), Novosibirsk Oblast (53.730N, 77.866E).

46 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

notes give good evidence to consider the interpretation secure. The slides were
then clear enough to score chiasma localization in the acrocentric autosomes (in 14
nuclei with at least ten autosomes scorable, the numbers of proximal vs. interstitial
vs. terminal chiasmata were 76:12:50 respectively). Five of these nuclei showed a
clear count of 14 acrocentric autosomes. The sex chomosomes were clear in several
nuclei, consisting of a “V” shaped trivalent with a metacentric at the point of the
“V”, to each arm of which was paired an acrocentric. One of those acrocentrics was
decondensed (heteropycnotic) in its centromeric half, and lying alongside it achi-
asmately was a decondensed acrocentric, thus in total making a figure of four. The
achiasmate pairing and heteropycnosis suggest those acrocentrics have ancestral
X material, as in the XXXY Habronattus (Maddison 1982, Maddison & Leduc-
Robert 2013), which this resembles strongly. Three pairs of second division nuclei
showed one member with 15 acrocentrics, the other with 12 acrocentrics and a
metacentric. Together this points to an XaXaXaYm sex chromosome system.

Attulus (Sitticus) fasciger: 26a+XaXa0 (Fig. 154). Many clear nuclei show the classical
13 acrocentric bivalents and two acrocentric X’s (heterpycnotic, side by side or
apart) off toward one pole. A few division-2 nuclei are consistent with this (three
nuclei with 13 similar acrocentrics; one nucleus with 13 similar and two more
condensed acrocentrics).

Attulus (Sitticus) finschi: 28a+XaXa0, with one autosome much smaller. This score re-
lies primarily on old notes, which indicate 28 acrocentric autosomes, one much
smaller than the others, and two acrocentric Xs. From the Chinook Lake speci-
men we have been able to re-score eight nuclei in first division with 15 figures, all
appearing as acrocentrics, and one much smaller than the others. The quality of
those nuclei is now too poor to distinguish the Xs. However, three other metaphase
nuclei in which the autosomes are not countable show clearly the two acrocentric
Xs heteropycnotic and lying side by side and toward one pole.

Attulus (Sitticus) pubescens: 26a+XaXmYa (Figs 155-163). Many nuclei indicate 26
acrocentric autosomes, but relatively few show the sex chromosomes clearly, ei-
ther because they are folded over themselves, or the X, is not clearly associated
with the others. However, many first division nuclei show a peculiar figure with a
metacentric (X,) whose shorter arm is paired terminally with an acrocentric (Y).
The longer arm of the X, is heteropycnotic, and is occasionally seen with the X,
lying achiasmately beside it. This behaviour suggests that the metacentric and loose
acrocentric are X’s, and this is supported by two cases of paired second division
nuclei: in each, one of the pair shows 14 all-acrocentric chromosomes, while its
partner shows more than 14 chromosomes, two of which are heteropycnotic. All
though the latter were not fully countable, in total 24 second division nuclei were
countable, 12 with 14 acrocentrics, and 12 with 14 acrocentrics plus a metacen-
tric. Together these point to one metacentric and one acrocentric X going to one
pole, in addition to 13 acrocentric autosomes, and one acrocentric Y to the other.

Attulus (Sitticus) terebratus: 26a+XaXa0 (Fig. 164). Several well-spread first meta-
phase show the two acrocentric Xs side by side and off to one pole, accompa-

Sitticine jumping spiders 47

4
|
C) (S) (CO) (Attinella dorsata)
@
O)

& . S @ Aittinetia concolor
0 =S O Jollas cupreus
SN. @ Sittisax ranieri
® SYX<Z @ Sittisax saxicola
= O Tomis manabita
S @ Aittutus finscni
Ex @ Attulus pubescens
i Y OO = O Aittulus fasciger
? - O => O© Aittulus terebratus
0 Oo => O Attulus ammophilus
0 OO => (Attulus burjaticus)
Se @ Attulus cutleri
> od @ Aittulus striatus
> C = O Aittulus inexpectus
C) = O Aittulus caricis
o » (ae )(@) (Attulus floricola/rupicola)
165 O => @ Aittulus floricola

Figure 165. Chromosome evolution in sitticines. Ancestral nodes show the most parsimonious recon-
struction of the evolution of Y via X-autosome fusions (black) from the X,X,0 sex chromosome system
(white). Phylogeny from Figure 48 with species added as follows: Attinella concolor is very similar in body
and genitalia to A. dorsata; likewise Sittisax saxicola to S. ranieri; Attulus caricis position based on COI results
(Fig. 96). The similar pair A. cudleri and A. striatus were placed as sisters to the floricola group based on their
inclusion in the floricola group by Logunov and Kronestedt (1997) and in Sittiflor by Prészynski (2017a).
Base chromosome number is directly the number of autosomes if the species has XXO sex chromosomes, but
is interpreted as the number of autosomes +2 if the species has XXY sex chromosomes (apparently derived
by a single fusion that would have consumed an autosomal pair), or + 4 if XXXY (apparently derived by

two fusions that would have consumed two pairs). Uncertain scoring is shown by parentheses (see Table 2).

nied by 13 pairs of acrocentric autosomes. Two second division nuclei show 15
acrocentrics, two of which are especially condensed (thus, the Xs), while one
shows 13 normal acrocentrics.

48 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

Chromosome evolution

While salticids are fairly conservative in basic chromosome complement, with most
species showing 26 acrocentric autosomes and X,X,0 sex chromosomes (Maddison
1982; Araujo et al. 2016, Araujo et al. 2019), sitticines are striking for their diver-
sity. Ihe distribution of chromosome complements on the reconstructed phylogeny
(Fig. 165) suggests that neo-Y chromosomes arose four separate times; the alternative,
assuming a Y was ancestral, is much less parsimonious, requiring seven losses to XXO.
Outgroups also favour XX0 as ancestral in sitticines: it is very much the most common
sex chromosome system in salticids, and the alternatives are phylogenetically scattered,
with no known Y chromosomes in other amycoids (Maddison 1982; Araujo et al.
2016, Araujo et al. 2019). Four X-autosome fusions among 18 species represents a
phylogenetic density approximately as high as in Habronattus (Maddison and Leduc-
Robert 2013), but the resulting forms of sex chromosomes are more varied in Sitticus.
The ancestral autosome number in sitticines is unclear. Among the species with
XX0, some have 26 autosomes, others have 28. Assessing a comparable autosome
number with neo-Y species requires interpretation, as the neo-Y system itself binds one
or more autosomal pairs with the X chromosomes, as indicated in part by distinctive
condensation patterns. If (as in Habronattus, Maddison and Leduc-Robert 2013) we
interpret the XXY systems as having one pair of autosomes bound into the sex chromo-
somes, and XXXY as having two pairs, then (for example) the 26a+XaXaYa of A. cutleri
is interpreted as having a base number of 28 (26 free and two bound). The rightmost
(red and white) column of Fig. 165 shows these interpreted base numbers. The most
parsimonious interpretation would then consider that red (28) is ancestral for the en-
tire clade of Attulus, reverting back to the typical salticid number (26) multiple times.
The ancestral node of the Jollas- Tomis clade, and the root of the Sitticini, could be 26 or
28 equally parsimoniously if the expected outgroup condition of 26 were not imposed.
An unanticipated but consistent correlation between base autosome number and
the presence of neo-Y is seen in Fig. 165, regardless of how we interpret the ancestral
state for base autosome number. The pattern is phylogenetically repeated: each of the
four separate neo-Y origins occurs in a 28-autosome lineage, and for each the closest
lineage with 26 has XX0. We have no suggestion as to why there might be such a cor-
relation. This pattern is unlikely to be a tautological consequence of our counting rule
that interprets XXY/XXXY systems as incorporating two/four autosomes. The counting
rule is derived (partially) independently, from condensation patterns and meiotic ori-
entation. Even lacking an independent argument within sitticines, we could import the
counting rule from Habronattus, where such an interpretation is well supported by mei-
otic behaviour and chromosome counts (Maddison 1982, Maddison and Leduc-Robert
2013). We do not know how to explain a correlation between an extra pair of autosomes
and the presence of neo-Y, but it is perhaps relevant that in all of the 28a+XaXa0 spe-
cies, one of the chromosome pairs is especially small, half or less the size of the others.
If these small chromosomes are supernumerary (B) chromosomes, it is possible
that there is considerably more variation within species than our small sample sizes can

Sitticine jumping spiders 49

detect. Undetected intraspecific variation in autosomes or sex chromosomes would
not negate our basic evolutionary conclusions. Were we to find species variable with
respect to the presence of a neo-Y chromosome, for example, it would point to even
more transitions between XX0O and XXY/XXXY.

Our uncertainty about chromosome complement in some species does not strong-
ly affect our conclusions about homoplasy or correlations, though it could affect a
detailed reconstruction of the evolution of autosome number, or of particular fusions
involved in a neo-Y system. For instance, if we delete autosome number for Attinella
dorsata and Attulus burjaticus (the two species with uncertain counts) from Fig. 165,
the ancestral states reconstructed by parsimony become ambiguously 28 or 26. Al-
though we are uncertain about the detailed intepretation of sex chromosomes in A.
rupicola/floricola and Sittisax saxicola, we conclude that they do have Y chromosomes,
and thus the reconstruction of Y chromosome evolution is not affected. The scope of
uncertainty allows one possible contradiction to our assessments above: should we
be incorrect about the autosome count of A. rupicola/floricola, this may be a species
in which a Y chromosome arose in the context of only 26 autosomes. Otherwise, the
ambiguities do not change the interpretation of a correlation between a base number
of 28 autosomes and neo-Y.

Chromosome evolution of sitticines will not be well understood, however, until a
larger sample of species and specimens is obtained, given the high diversity seen in our
small sample. Our data hint to the possibility of rapid evolution provoked by special
mechanisms.

Acknowledgements

We are grateful to several colleagues who made special efforts to provide us access to
specimens: to Galina Azarkina for greatly facilitating collecting in Siberia, to Gergin
Blagoev for preparing and taking photographs of Attulus sylvestris specimens used for
barcoding, and to Cristian Grismado for taking photographs of the holotype of Jollas
puntalara Galiano. We thank Petra Kranebitter of the Naturmuseum Siidtirol/ Museo
di Scienze Naturali dell’Alto Adige for lending us the specimen of Attulus (Sittilong)
longipes, Simone Ballini for collecting it, and Tobias Bauer for leading us to it. Laura
Leibensperger and Jennifer Zaspel both helped generously in our attempts to trace
what are the true types of Attus palustris Peckham and Peckham. Laura Leibensper-
ger and Gonzalo Giribet of the MCZ, and Lou Sorkin and Lorenzo Prendini of the
AMNH provided loans of important type material. We thank Gonzalo Giribet for
kindly providing lab space and resources to sequence Attulus longipes. Junxia Zhang
suggested euophryine identifications for two species misplaced in Sitticus. Helpful
comments on the manuscript were provided by G.B. Edwards, D. Logunoy, J. Miller,
and two anonymous reviewers. We especially thank one of the anonymous reviewers
for provoking us to examine our chromosome data more thoroughly. Funding was

provided to WPM through an NSERC postgraduate scholarship and an NSERC Dis-

50 Wayne P Maddison et al. / ZooKeys 925: 1-54 (2020)

covery grant, and to DRM by the Harold E. and Leona M. Rice Endowment Fund
at Oregon State University. Data collection in the Hedin lab was supported by US
National Science Foundation (DEB 1754591).

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